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FLORA OF THE USSR 
Volume XXIII 


Bignoniaceae — Valerianaceae 


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FLORA OF THE USSR 


Initiated under the supervision and chief editorship 
of Academician V.L. Komarov 


VOLUME XXIII 


Bignoniaceae—Valerianaceae 


/ a . ’ 
Volume Editor ( i j } 
B.K. SCHISCHKIN ™ 


General Scientific Editors 


Stanwyn G. Shetler 
Galina N. Fet 
Sergei L. Mosyakin 


Translated from Russian 


Smithsonian Institution Libraries 
Washington, D.C. 
2000 


SMIN B87-104 


Flora SSSR, Tom XXIII 
Akademiya Nauk SSSR Publishers, 
Moscow-Leningrad, 1955 


© 1999 Amerind Publishing Co. Pvt. Ltd., New Delhi 
Translator & Editor: V. Pandit 


Library of Congress Cataloging-in-Publication Data 
Flora SSSR. English 
Flora of the U.S.S.R. = Flora SSSR. 


Cover title: Flora U.S.S.R. 

Translation of: Flora SSSR. 

At head of title, v. —23. V.L. Komarov Botanical 
Institute. Academy of Sciences of the USSR. 

Reprint. Originally published: Leningrad - 
Izdatel'stvo Akademi nauk SSSR, 1934— 

Vols. Have imprint: Washington, D.C. : Smithsonian 
Institution Libraries and the National Science 
Foundation 

Includes bibliographies and indexes. 


I. Botany—Soviet Union—Classification. 
2. Plants—Identification. I. Komarov, V. L. 
(Vladimir Leontevich), 1869-1945. II. Botanicheskii 
institut im. V.L. Komarova. III. Title: Flora SSSR. 
III. Title: Flora U.S.S.R. V. Title. 
K321.F69613 1985 85-904577 


Translated and published under an agreement with the Smithsonian Institution Libraries, 
Washington, D.C., by Amerind Publishing Co., 66 Janpath, New Delhi 110 001. 


Printed at Baba Barkha Nath Printers, 26/7 Najafgarh Road Industrial Area, 
New Delhi 110015 


INSTITUTUM BOTANICUM NOMINE V.L. KOMAROVII ACADEMIAE SCIENTIARUM URSS 


FLORA 
URSS 


(FLORA UNIONIS RERUMPUBLICARUM 
SOCIALISTICARUM SOVIETICARUM) 


XXIII 


EDITIO ACADEMIAE SCIENTIARUM URSS 
MOSQUA 1958 LENINGRAD 


V.L. KOMAROV BOTANICAL INSTITUTE 
ACADEMY OF SCIENCES OF THE USSR 


Contributors 


E.G. Bobrov, I.T. Vassilczenko, S.G. Gorschkova, 
Yu.S. Grigorev, V.I. Grubov, P.I. Dorofeev, I.A. Iljinskaya, 
M.V. Klokov, L.A. Kuprianova, I.A. Linczevsky, 
I.V. Novopokrovsky, E.G. Pobedimova, M.G. Popov, 
A.I. Pojarkova, B.K. Schischkin, E.I. Steinberg, N.N. Tzvelev 


SCIENTIFIC EDITOR’S PREFACE TO VOLUME XXIII 


For practical reasons, I have concentrated my editorial review of 
this volume on the discussions and the habitat and distribution 
statements. Dr. Fet also has reviewed the entire volume and paid 
particular attention to the geographic and place-name terminology in 
the distribution statements following the morphological descriptions. 
Dr. Mosyakin, during a brief visit to the Smithsonian Institution, was 
able to review more than half of the manuscript, and his comments 
have been enormously helpful. We are confident of the general accuracy 
of the translation but also recognize that there will be imperfections. 


September 1999 


STANWYN G. SHETLER 

Curator of Botany Emeritus 
National Museum of Natural History 
Smithsonian Institution 

Washington, DC 20560 


Assisted by Galina N. Fet 

Huntington, West Virginia 25701 

and 

Sergei L. Mosyakin 

Head of Vascular Plants Department 

N.G. Kholodny Institute of Botany 
National Academy of Sciences of Ukraine 
Kiev, GSP-1, 252601 Ukraine 


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PREFACE 


The twenty-third volume of the “Flora of the USSR,” published 
after the twenty-fourth volume, contains the descriptions of the species 
belonging to the families Bignoniaceae to Valerianaceae. 

The family Orobanchaceae, initially treated by the late 
I.V. Novopokrovsky, was revised by N.N. Tzvelev, at the request of 
the editor. 

Much new work has been done on the taxonomy of the family 
Rubiaceae, with the collaboration of M.V. Klokov, I.A. Linczevsky, 
E.G. Pobedimova, A.I. Pojarkova and B.K. Schischkin. Especially much 
new work has been done by E.G. Pobedimova in studying the species 
of bedstraw and woodruff. 

The large honeysuckle family, Caprifoliaceae, has been thoroughly 
revised by A.I. Pojarkova. Here, she presents a new taxonomic 
treatment of the genera Lonicera and Sambucus. 

The genus Valeriana has also been newly revised. The author, 
V.I. Grubov, considered it necessary to lump many of the species 
described from the USSR. Grubov documented his own views earlier 
(1955) in a separate paper. 


Editors. 


777 


CONTRIBUTORS 


Families Bignoniaceae and Pedaliaceae 

Family Martyniaceae 

Family Orobanchaceae (excluding genus 
Mannagettaea) 

Genus Mannagettaea 

Family Lentibulariaceae 

Family Globulariaceae 

Family Phrymaceae 

Family Plantaginaceae 

Characters of family Rubiaceae, genera 
Karamyschewia, Gaillonia, Sherardia, 
Crucianella, Phuopsis, Valantia, 
Valerianella, Pseudobetckea 

Genus Asperula (excluding sections 
Cynanchica, Sherardiana and 
Oppositifoliae), genera Galium and 
Microphysa 

Section Cynanchica of genus Asperula 

Sections Sherardiana and Oppositifoliae 
of genus Asperula, genera Leptumis, 
Mitchella, Callipeltis, and family 
Adoxaceae 

Genus Rubia and family Caprifoliaceae 

Characters of family Valerianaceae, 
Key for characteristics of genera and 
genus Valeriana 

Genera Patrinia and Kentranthus 

Index to names of fossil plants 


: LT. Vassilczenko 
: S.G. Gorschkova 
: I.V. Novopokrovsky 


: M.G. Popov 

: E.I. Steinberg 

: E.G. Bobrov 

: L.A. Kuprianova 
: Yu.S. Grigorev 
: LA. Linczevsky 


: B.G. Pobedimova 


: M.V. Klokov 
: B.K. Schischkin 


: A.I. Pojarkova 
: V.I. Grubov 


: LA. Iliaskaya 
: P.I. Dorofeev 


Plates prepared by: N.A. Moiseeva, I-II; VI-—X XIII, XXX—XXXII;, 
E.S. Gaskevich—III, IV, XX XIII, XXXIV; M.R. Gabe—V, VI, XXIV- 
XXVII; A.E. Likas—XXVII, Z.V. Kobyletskaya-XXIX 


CONTENTS 


SCIENTIFIC EDITOR’S PREFACE TO Be Vil 
VOLUME XXIll 

PREFACE as ix 

CONTRIBUTORS we X 

SYSTEMATIC INDEX OF SPECIES OF THE sae XV 


TWENTY-THIRD VOLUME OF “FLORA OF THE 
USSR” 


FAMILY CXLIT. BIGNUNIACEAE PERS. 1 
Genus *Campsis Lour. 1 
Genus 1367. Incarvillea Juss. io 3 
Genus 1368. Niedzwedzkia B. Fedtsch. 5 
Genus *Catalpa Scop. i 7 

FAMILY CXLIV. PEDALIACEAE LINDL. ces 11 
Genus *Sesamum L. ay 11 
Genus 1369. Trapella Oliv. = 13 

FAMILY CXLV. MARTYNIACEAE LINK Ln 16 
Genus 1370. Proboscidea Moench Cos 16 

FAMILY CXLVI. OROBANCHACEAE LINDL. ni 17 
Genus 1371. Cistanche Hoffmg. and Link oe 19 
Genus 1372. Phelypaea Desf. me 26 
Genus 1373. Orobanche L. eh 28 
Genus 1374. Boschniakia C.A.M. mi 108 
Genus 1375. Mannagettaea H. Smith bei 109 


Genus 1376. Phacellanthus Sieb. et Zucc. ae 110 


Xli 


FAMILY CXLVII. LENTIBULARIACEAE LINDL. _... 111 


Genus 1377. Pinguicula L. ae 112 
Genus 1378. Utricularia L. ies 117 
FAMILY CXLVIII. GLOBULARIACEAE LINK = 123 
Genus 1379. Globularia L. a 123 
FAMILY CXLIX. PHRYMACEAE SCHAUER a: 126 
Genus 1380. Phryma L. Sab 126 
ORDER 37. PLANTAGINALES ENGL. za) 128 
FAMILY CL. PLANTAGINACEAE LINDL. - 129 
Genus 1381. Plantago L. 2 129 
Subgenus 1. Euplantago Harms os 132 
Subgenus 2. Psyllium (Juss.) Harms ae 154 
Genus 1382. Littorella Bergius 4. 156 
ORDER 38. RUBIALES ENGL. 3 157 
FAMILY CLI. RUBIACEAE JUSS. uel 157 
Subfamily 1. Cinchonoideae K. Schum. a 160 
Genus 1383. Karamyschewia Fisch. et Mey. cm 160 
Subfamily 2. Coffeoideae K. Schum. ” 161 
Tribe 1. Anthospermeae Cham. and Schlechtd. ie 161 
Genus 1384. Mitchella L. my 161 
Tribe 2. Spermacoceae Spreng. me 162 
Genus 1385. Gaillonia A. Rich. ms 162 
Tribe 3. Galieae Dumort. ae 166 
Genus 1386. Sherardia L. oe 166 
Genus 1387. Crucianella L. me 169 
Genus 1388. Phuopsis (Griseb.) Hook. f. ™ 183 
Genus 1389. Asperula L. ai 184 
Genus 1390. Leptunis Stev. sa 270 
Genus 1391. Microphysa Schrenk sigs 271 
Genus 1392. Galium L. ” Dae 
Genus 1393. Valantia L. emend. DC. si 362 


Genus 1394. Madder—Rubia L. bis 363 


Genus 1395. Callipeltis Stev. 
FAMILY CLII. CAPRIFOLIACEAE VENT. 
Tribe 1. Sambuceae (Kunth) DC. 
Genus 1396. Sambucus L. 
Tribe 2. Viburneae (Spach) Fritsch 
Genus 1397. Viburnum L. 
Genus 1398. Triosteum L. 
Tribe 3. Linnaeeae Fritsch 
Genus 1399. Abelia R. Br. 
Genus 1400. Linnaea Gronov. ex L. 
Tribe 4. Lonicereae R. Brown 
Genus 1401. Lonicera L. 
Subgenus 1. Chamaecerasus Rehd. 
Subgenus 2. Periclymenum Rehd. 
Tribe 5. Diervilleae C.A.M. 
Genus 1402. Weigela Thunb. 
FAMILY CLIII. ADOXACEAE FRITSCH 
Genus 1403. Adoxa L. 
FAMILY CLIV. VALERIANACEAE DC. 
Tribe 1. Patrineae Hoeck 
Genus 1404. Patrinia Juss. 
Tribe 2. Valerianeae (Hoeck) Graebn. 
Genus 1405. Valeriana L. 
Subgenus 1. Valeriana 
Genus 1406. Centranthus Neck. 
Genus 1407. Valerianella Mill. 


Genus 1408. Pseudobetckea (Hoeck) Lincz. 


ADDENDA XXII. DIAGNOSES PLANTARUM 
NOVARUM IN TOMO XXIII FLORAE URSS 
COMMEMORATARUM 


INDEX ALPHABETICUS 


Xill 


396 
397 
399 
400 
421 
421 
437 
439 
439 
443 
446 
446 
456 
544 
549 
549 
558 
558 
560 
561 
561 
566 
567 
572 
609 
611 
649 


651 
711 


3 ‘Poms 1084 atitohelta: bi | 


eee 


2. . Speneancoront Sprea, seut sins 2081 


vi SYSTEMATIC INDEX OF SPECIES OF VOL. XXIII 
OF THE “FLORA OF THE USSR” 


Family CXLIII. Bignoniaceae Pers. 


Genus* Campsis Lout. 


Page no.* 
1. C. radicans (L) Seem Riss 2 
Genus 1367. Incarvillea Juss. 

13189 1. FL. olgae Regi. bis 3 
13190 2. O. sinensis Lam. phe 4 
Genus 1368. Niedzwedzkia B. Fedtsch. 

1. N. semiretschenskia B. Fedtsch. bee = 
Genus* Catalpa Scop. 
1. C. bignonioides Walt. oe 10 
2. C. speciosa Ward. oF 10 
3. C. ovata G. Don Wis 11 
Family CXLIV. Pedaliaceae Lindl. 
Genus* Sesamum L. 
1. S. orientale L. B 12 
Genus 1369. Trapella Oliv. 
1. T. sinensis Oliv. ie 14 


Family CXLV. Martyniaceae Link 
Genus 1370. Proboscidea Moench 


tReproduced from the Russian original. Russian page numbers appear in the left- 
hand margin of the text—General Editor. 


Vill 


XVI 


13200 


13210 


1. P. louisiana (Mill.) Woot et Stand. 


ON 


7. 


a) ac) ac 


WR WN ES 


= =| =| SS SS 
ARONESSMNANAWNE 


ae. 


C: 
Re Se 
(6s 
C: 
C. 


C. 


SISss919 SS 9 9S So 9° °°.°0'")'9 


Family CXLVI. Orobanchaceae Lindl. 
Genus 1371. Cistanche Hoffmg. et Link 


Section 1. Eucistanche G. Beck 


tubulosa (Schenk) R. Wight 
flava (C.A.M.) Korsh. 
mongolica G. Beck 
salsa (C.A.M.) G. Beck 
ambigua (Bge.) G. Beck 
Section 2. Heterocalyx G. Beck 


fissa (C.A.M.) G. Beck 


Section 3. Cistanchiella G. Beck 


ridgewayna Aitch. et Hems1. 


Genus 1372. Phelypaea Desf. 


. coccinea (M.B.) Poir. 
. helenae Popl. 
. tournefortii Desf. 


Genus 1373. Orobanche L. 
Section 1. Trionychon Wallroth 


Subsection 1. Holoclada Novopokr. 


. purpurea Jacq. 

. uralensis G. Beck 

. caesia Rchb. 

. pulchella (C.A.M.) Novopokr. 


bungeana G. Beck 


. iberica (G. Beck) Tzvel. 
. arenaria Borkh. 


androssovii Novopokr. 


. karatavica Pavl. 
. coelestis Boiss. et Reut. 


borissovae Novopokr. 


. kelleri Novopokr. 

. cilicica G. Beck 

. septemloba (G. Beck) Tzvel. 
. serratocalyx G. Beck 


18 


23 
23 
24 
24 
25 


26 


2h 


13220 


13230 


13240 


13250 


ee Bee) (Ss) (Ss) 1S) Stes) 


SOS SSS GSS Gia) Ga GS Seeley GS GG SiS 


Subsection 2. Pleioclada Novopokr. 


. hirtiflora (Reut.) Tzvel. 

. aegyptiaca Pers. 

. mutelii F. Schultz 

. brassicae Novopoktr. 

. hohenackeri (Reut.) Tzvel. 
. brachypoda Novopokr. 

. oxyloba (Reut.) G. Beck 

. orientalis G. Beck 

. sogdiana Novopokr. 

. dalmatica (G. Beck) Tzvel. 


nana Noé 
ramosa L. 


Subsection 3. Inflatae G. Beck 


. coerulescens Steph. 

. korshinskyi Novopokr. 
. glaucantha Trautv. 

. pycnotachya Hance 


amurensis (G. Beck) Kom. 
amoena C.A.M. 

ariana Gontsch. 

hansii Kerner 


. clarzei Hook. f. 
. sintenisii G. Beck 
. solenanthi Novopokr. et Pissjauk. 


sulphurea Gontsch. 
cernua Loefl. 
cumana Wallr. 
brevidens Novopokr. 


. solmsii Clarke 


badchysensis Novopokr. et V. Nikitin 
camptolepis Boiss. et Reut. 
kotschyi Reut. 


. gigantea (G. Beck) Gontsch. 
. sordida C.A.M. 

. vitellina Novopokr. 

. glabricaulis Tzvel. 

. gamosepala Reut. 

. lutea Baumg. 

. linczevskyi Novopokr. 

. teucrii Holand. 


XVil 


XVIill 


55. O. schelkovnikovii Tzvel. of 90 
56. O. vulgaris Poir. ace 91 
57. O. hymenocalyx Reut. ae 92 
58. O. alba Steph. SH 93 
13260 59. O. pallidiflora Wimm. et Grab. re 94 
60. O. reticulata Wallr. aN 95 
61. O. grigorjevii Novopokr. he 95 
62. O. raddeana G. Beck ths 96 
63. O. transcaucasica Tzvel. bf 97 
64. O. owerinii G. Beck re, 97 
65. O. crenata Forsk. aa 98 
66. O. picridis F. Schultz a 101 
67. O. minor Sutt. oe 102 
68. O. versicolor F. Schultz was 103 
13270 69. O. hederae Duby a6 104 
70. O. connata C. Koch dict 104 
71. O. armena Tzvel. oa 105 
72. O. kurdica Boiss. et Hausskn. ia 105 
73. O. Grossheimii Novopokr. ius 106 
74. O. major L. sas 107 
75. O. flava Mart. de 108 
76. O. inulae Novopokr. et Abram. sad 108 
77. O. krylowii G. Beck af 109 
78. O. alsatica Kirschl. Be 110 
79. O. rosea Tzvel. i 111 
13280 80. O. gracilis Smith a 112 
81. O. colorata C. Koch st 113 


Genus 1374. Boschniakia C.A.M. 
1. O. rossica (Chem. et Schlechtend.) ai 114 
B. Fedtsch. 
Genus 1375. Mannagettaea H. Smith 
1. O. ircutensis M. Pop. te 115 


Genus 1376. Phacellanthus Sieb. et Zucc. 
1. P. tubiflorus Sieb. et Zucc. of 116 


13290 


13300 


Family CXLVII. Lentibulariaceae Lindl. 


Tribe Utricularieae Kaminski 
Genus 1377. Pinguicula L. 


. variegata Turcz. 

alpina L. 

. villosa L. 

. bicolor Woloszck. 

. vulgaris L. 

glandulosa Trautv. et Mey. 
. spathulata Ldb. 


Pips Da Ale atte Dal 
a Da- Jia “ia -Iia Tati 


Genus 1378. Utricularia L. 


. intermedia Hayne 
minor L. 

bremii Heer 

. Japonica Makino 
. major Schmidel 

. vulgaris L. 


ANWRWNH 
Se cee ee 


Family CXLVIII. Globulariaceae Link 
Genus 1379. Globularia L. 
1. G. aphyllanthes Cr. 
2. G. trichosantha Fisch. et Mey. 
Family CXLIX Phrymaceae Schauer 
Genus 1380. Phryma 1. 
1. Ph. leptostachya:L. 


Family CL. Plantaginaceae Lindl. 
Genus 1381. Plantago L 
Subgenus 1. Euplantago Harms 
Section 1. Polyneuron Decne. 


1. P. major L. 


Section 2. Micropsyllium Decne. 


2. P. tenuiflora Waldst. et Kit. 
3. P. polisperma Kar. et Kir. 


xix 


119 
119 
120 
120 
121 
122 
122 


124 
125 
125 
126 
127 
127 


129 
130 


132 


138 


139 
140 


xi 


XX 


13310 


13320 


7. 


18. 


19. 


ach ae) 


se) acl gel ac) ae) 


7 Sw 0 


aa Iai ia a 


Section 3. Palaeopsyllium Pilger 


. cornuti Gouan 


Section 4. Holopsyllium Pilger 


. macrocarpa Cham. et Schlechtd. 


Section 5. Coronopus DC. 


. coronopus L. 
. maritima L. 


Section 6. Mesembrinia Decne. 


. schwarzenbergiana Schur 
. depressa Willd. 

. camtschatica Link 

. arachnoidea Schrenk 

. komarovii Pavl. 


Section 7. Lamprosantha Decne. 


media L. 


. Stepposa Kuprt. 
. maxima Juss. 
. canescens Adams. 


Section 8. Oreades Decne. 


. atrata Hoppe 


Section 9. Gentianoides Pilger 


. gentianoides Sibth. et Smith 


Section 10. Arnoglossum Decne. 


. lanceolata L. 


Section 11. Leucopsyllium Decne. 


. ovata Forsk. 
. minuta Pall. 


loeflingii L. 
notata Lagasca 


. lachnantha Bge. 
. lagocephala Bge. 


141 


142 


142 
143 


145 
145 
146 
146 
147 


148 
148 
151 
152 


152 


155 


156 


loi) 
158 
159 
159 
160 
160 


26. 
2H. 


xii 


13350" 1 


kk WN = 


—" 


Subgenus 2. Psyllium (Juss.) Harms 
Section 12. Psyllium (Juss.) Barneoud 
P. indica L. 
P. psyllium L. 


Genus 1382. Litorella Bergius 


. L. uniflora (L.) Aschers. 


Subseries 38. Rubiales Engl. 
Family CLI. Rubiaceae Juss. 
Subfamily I. Cinchonoideae K. Schum. 
Tribe 1. Oldenlandieae K. Schum. 


Genus 1383. Karamyschewia Fisch. et Mey. 


K. hedyotoides Fisch. et Mey. 


Subfamily II. Coffeoideae K. Schum. 


Tribe 1. Anthospermeae Cham. et Schlechtd. 


Genus 1384. Mitchella L. 


. M. undulata Sieb. et Zucc. 


Tribe 2. Spermacocoeae Spreng. 


Genus 1385. Gaillonia A. Rich. 


Section 1. Microstephus (Jaub et Sp.) Boiss. 


. G.-bucharica B. Fedtsch. et Desjat. 
. G. szovitsii DC. 

. G. bruguierei A. Rich. 

. G. asperuliformis Lincz. 


Tribe 3. Galieae Dumort. 


Genus 1386. Sherardia L. 


. S. arvensis L. 


Genus 1387. Crucianella L. 


. C. schischkinii Lincz. 


2. C. buchrica B. Fedtsch. 


XXi 


161 
162 


163 


168 


169 


170 
171 
171 
175 


176 


178 
179 


Xill 


XXii 


13340 


13350 


13360 


1. 


10. 
11. 


12. 


13. 
14. 
15. 
16. 
Ne 


SON AO ee 
> Dd SSD DB DS D 


Genus 1388. Phuopsis (Griseb.) Hook. f. 
P. 


> > 


> > > > D> 


C. 
i Ge 
. divaricata Korov. 


OOO) Gee @ @ 


sabulosa Korov. et Krasch. 
filifolia Rg]. et Winki. 


baldshuanica Krasch. 


. exasperata Fisch. et Mey. 

. chlorostachys Fisch. et Mey. 
. angustifolia L. 

. latifolia L. 

. gilanica Trin. 

. suaveolens C.A.M. 

. Sintenisii Bornm. 


Stylosa (Trin.) Hook. f. 


Genus 1389. Asperula L. 


Section 1. Cruciana Gris. 


. mollugionoides (M.B.) Rchb. 
. prostrata (Adams) C. Koch. 
. albovii Mand. 

. glomerata (M.B.) Griseb. 

. hirsutiuscula Pobed. 

. ferganica Pobed. 

. pamirica Pobed. 

. balchanica Bobr. 

. turcomanica Pobed. 


Section 2: Monocephala Pobed. 


. caucasica Pobed. 
. propinqua Pobed. 


Section 3. Trichocarpae Pobed. 


. odorata L. 


Section 4. Cynanchica DC. 


. cemiamicta Klok. 

. cynanchica L. 

. creticola Klok. 

. tephrocarpa Czern. 

. exasperata V. Krecz. 


180 
183 
184 
185 
185 
186 
187 
188 
189 
190 
191 


192 


205 
206 
207 
208 
211 
212 
212 
213 
214 


215 
216 


217 


223 
224 
225 
226 
227 


13370 


13380. 


13390 


xiv 13400 


56. 
a7. 


>> BRD DDD DDS DD R.DS RDS. SPR. RR. RRR RdD DD DDD DD DD>D>DD 


> > 


. petraea V. Krecz. 
. supina M.B. 

. caespitans Juz. 

. tranzshelii Klok. 
. stevenii V. Krecz. 


bidentata Klok. 


. hypanica Klok. 
. attenuata Klok. 
. rumelica Boiss. 


craniticola Klok. 

markothensis Klok. 
lipskyana V. Krecz. 
aemulans V. Krecz. 


. praevestita Klok. 

. vestita V. Krecz. 

. infracta Klok. 

. praepilosa V. Krecz. 

. cimmerica V. Krecz. 

. biebersteinii V. Krecz. 

. pedicellata Klok. 

. dasyantha Klok. 

. accrescens Klok. 

. cristata (Somm. et Lev.) V. Krecz. 


alpina M.B. 


. intersita Klok. 
. abchasica V. Krecz. 


pontica Boiss. 
affinis Boiss. et Huet 


. dolichophylla Klok. 


woronowii V. Krecz. 
kemulariae Manden. 


. setulosa Boiss. 

. savranica Klok. 

. graveolens M.B. 

. leiograveolens M. Pop. et Chrshan. 
. diminuta Klok. 

. laevissima Klok. 

. danilewskiana Basin. 


Section 5. Galioideae DC. 


. octonaria Klok. 
. campanulata (Vill.). Klok. 


XXill 


228 
229 
230 
231 
232 
233 
235 
235 
236 
239 
240 
241 
241 
242 
243 
244 
245 
246 
247 
247 
249 
249 
250 
253 
254 
255 
256 
257 
258 
259 
259 
260 
261 
262 
262 
263 
264 
264 


265 
266 


XXIV 


13410 


13420 


13430 


AD 


1. 


> > > D> D> 


> > 


> > b> 


oe se 


. tyraica Bess. 

. galioides M.B. 
. xerotica Klok. 

. paniculata Bge. 


kryloviana Serg. 


. tinctoria L. 


platygalium Maxim. 
maximowiczii Kom. 


. rivalis Sibth. et Sm. 
. aparine M.B. 
. karataviensis Pavl. 


Section 6. Brachyanthae Pobed. 


. humifusa (M.B.) Bess. 
. insuavis Pobed. 


Section 7. Oppositifoliae Schischk. 


. oppositifolia Rgl. et Schmalh. 
. popovii Schischk. 
. laevis Schischk. 


Section 8. Sherardiana DC. 


. arvensis L. 

. setosa Jaub. et Sp. 
. azurea Jaub. et Sp. 
. taurica Paczoski 

. gracilis C.A.M. 


Genus 1390. Leptunis Stev. 


. trichoides (J. Gay) Schischk. 


Genus 1391. Microphysa Schrenk 


M. elongata (Schrenk) Pobed. 


Genus 1392. Galium L. 
Section.1. Depauperata Pobed. 


Subsection 1. Quadrifolia Pobed.. 


. G. soongoricum Schrenk 


267 
267 
268 
271 
271 
272 
273 
273 
274 
275 
275 


276 
278 


278 
280 
280 


281 
282 
283 
284 
285 


285 


286 


299 


XV 


13440 


13450 


8. 
9. 
10. 
11. 
12. 
13 


QAAAADA Qa 


a0 


QAAAAAAAAAADAN 


Subsection oy Senifolia Pobed. 


. triflorum Michx. 
. trifloriforme Kom. 


Section 2. Aparine DC. 


Subsection 1. Camptopoda Boiss. 


. tricorne Stokes 
. linczevskyi Pobed. 


Subsection 2. Leucaparine Boiss. 


. aparine L. 
. spurium L. 


Section 3. Pseudoaparine Lange 


. tenuissimum M.B. 

. trichophorum Kar. et Kir. 

. transcaucasicum Stapf 

. chloreleucum Fisch. et Mey. 
. decaisnei Boiss. 

. eldaricum Grossh. 


Section 4. Asperae (Moench) DC. 


‘ verticillatum Danth. 
. apsheronicum Pobed. 
. nupercreatum M. Pop. 


Section 5. Cruciatae DC. 


. vernum Scop. 
. schischkinii Pobed. 
. krylovii Ijin 


elbrussicum Pobed. 


. cruciata (L.) Scop. 
. tauricum (Willd.) Roem. et Schult. 


sosnowskyi Manden. 


. braunii Zelen. 

. coronatum Sibth. et Sm. 

. kopetdaghense Pobed. 

. sevanense Pobed. 

. cordatum Roem. et Schult. 


XXV 


300 
303 


304 
305 


306 
306 


308 
308 
309 
310 
311 


3 


312 
313 
313 


314 
318 
318 
319 
319 
320 
321 
322 
322 
323 
324 
324 


XXVi 


13460 


13470 


13480 


13490 


Qa 


DQAAIAAAAYN 


Gah GY Gy ay G) GG) a) G) 


Gl) GiGy G) Gl G) GiiGyian @ 


. pedemontanum (Bell.) All. 
. pseudopolycarpon Somm. et Lev. 


Section 6. Cymogaliae Pobed. 


. paradoxum Maxim. 
. scabrum L. 
. kamtschaticum Steller 


Section 7. Aparinoides Jord. 


. palustre L. 

. Saturejifolium Trev. 
. karakulense Pobed. 
. elongatum Presl 

. krymense Pobed. 

. trifidum L. 

. ruprechtii Pobed. 

. baicalense Pobed. 


Section 8. Leptogalium Lange 


. hercynicum Weig. 
. tianschanicum M. Pop. 
. anisophyllum Vill. 


pumilum Murr. 
vartanii Grossh. 
kiapazi Manden. 
uliginosum L. 


. dahuricum Turcz. 
. vassilezenkoi Pobed. 
. pojarkovae Pobed. 


Section 9. Platygalia DC. 


. articulatum Lam. 
. rubioides L. 
. volgense Pobed. 


ussuriense Pobed. 
boreale L. 


. amurense Pobed. 

. amblyophyllum Schrenk 

. turkestanicum Pobed. 

. mugodsharicum Pobed. 

. Septentrionale Roem. et Schult. 


825 
325 


326 
329 
330 


S31 
332 
332 
233 
334 
334 
335 
336 


336 
337 
338 
338 
339 
340 
340 
341 
342 
345 


345 
347 
348 
348 
349 
350 
351 
S50 
352 
353 


XVii 


13500 


13510 


13520 


62. 


92. 
93. 


QQ 


QAAAAAAAARAAAAAA 


AAAAAIAAAAAAAAD 


. valantioides M.B. 


Section 10. Xanthogalia DC. 


. verum L. 
subuliferum Somm. et Lev. 


wirtgenii F. Schultz 


fistulosum Somm. et Lev. 
anfractum Somm. et Lev. 


majmechense Bordz. 
pamiroalaicum Pobed. 
aureum Visiani 
achurense Grossh. 


lacteum (Maxim.) Pobed. 


densiflorum Ldb. 
ruthenicum Willd. 


. Saurense Litw. 
. atropatanum Grossh. 
. kutzingii Boiss. et Buhse 


Section 11. Eugalium DC. 
. mollugo L. 


erectum Huds. 


. Juzepczukii Pobed. 


calcareum (Alb.) Pobed. 
schultesii Vest. 
polonicum Blocki 
caspicum Stev. 
grusinum Trautv. 
hyrcanicum C.A.M. 
mite Boiss. et Hoh. 


leiophyllum Boiss. et Hoh. 
. bullatum Lipsky 


grossheimii Pobed. 


. czerepanovii Pobed. 


Subsection. Coriacea Pobed. 


. coriaceum Bge. 
. brachyphyllum Roem. et Schult. 


Genus 1393. Valantia L. emend. DC. 


. muralis L. 


XXVIi 


354 


357 
358 
358 
359 
360 
361 
361 
362 
363 
363 
364 
367 
368 
369 
369 


370 
371 
Sie. 
32 
373 
374 
375 
375 
376 
377 
377 
378 
379 
379 


379 
380 


381 


XXVili 


13530 


13540 


XVill 


NO = 


~ 


= 
pe) 


16. 


By DD WW WW WW WWD 


Genus 1394. Rubia L. 


Section 1. Oligoneura Pojark. 


. cordifolia L. 
. chinensis Rgl. et Maack. 


Section 2. Meganthera Pojark. 


. tinctorum L. 
. lberica (Fisch. ex DC.) C. Koch 
. cretacea Pojark. 


Section 3. Chonanthe Pojark. 


. rezniczenkoana Litw. 


Section 4. Campylanthera Pojark. 


. deserticola Pojark. 

. chiralensis Ehrendorf. 
. tibetica Hook. f. 

. regelii Pojark. 

. komarovii Pojark. 

. dolichophylla Schrenk 


krascheninnikovii Pojark. 


. Jesoensis (Miq.) Miyabe et Miyake 
. tatarica (Trev.) Fr. Schmidt 


transcaucasica Grossh. 
laxiflora Gontsch. 
schugnanica B. Fedtsch. 


. rigidifolia Pojark. 


florida Boiss. 


. rechingeri Ehrerdorf. 


Genus 1395. Callipeltis Stev. 


cucullaris (Jusl.) Rothm. 


Family CLII. Caprifoliaceae Vent. 


Tribe 1. Sambuceae (Kunth) DC. 
Genus 1396. Sambucus L. 


Section 1. Eusambucus Spach 


. nigra L. 


387 
391 


393 
394 
396 


397 


399 
400 
401 
402 
403 
404 
406 
407 
408 
411 
412 
413 
414 
415 
416 


418 


424 


XX1X 


Section 2. Ebulus Spach 
2. S. ebulus L. bt 427 


Section 3. Olma (Raf.) Rehd. 


3. S. latipinna Nakai se? 429 
13550 4. S. coreana (Nakai) Kom. et Allis. be 430 
5. S. racemosa L. Ree 431 
6. S. tigrani N. Troitzky wee 433 
7. S. sibirica Nakai fe 434 
8. S. manshurica Kitag. Sa 436 
9. S$. kamtschatica E. Wolf na 437 
10. S. sachalinensis Pojark. ma 439 
11. S. sieboldiana (Miq.) Schwer. bs 441 
Tribe 2. Viburneae (Spach) Fritsch 
Genus 1397. Viburnum L. 
Section 1. Pseudotinus Clarke 
1. V. furcatum B1. se 445 
Section 2. Lantana Spach 
2. V> lantana L. au 447 
13560 3. V. burejaeticum Rg. et Herd. sa 448 
4. V. mongolicum (Pall.) Rehd. sie 450 
Section 3. Odontotinus Rehd. 
5..V. wrightii Miq. = 451 
6. V. orientale Pall. te 452 
Section 4. Opulus DC. 
7. V. sargenti Koehne Bi 454 
8. V. opulus L. ve 456 


Genus 1398. Triosteum L. 


1. T. sinuatum Maxim. an 458 


XXX 


Tribe 3. Linnaeeae Fritsch 
Genus 1399. Abelia R. Br. 
Section 1. Zabelia Rehd. 


Subsection 1. Corymbosae Rehd. 


1. A. corymbosa Rgl. et Schmalh. act 461 
xix Subsection 2. Biflorae Zabel 
2. A. coreana Nakai a 463 


Genus 1400. Linnaea Gronov. ex 'L. 


1. L. borealis L. igs 465 


Tribe 4. Lonicereae R. Brown 
Genus 1401. Lonicera L. 
Subgenus 1. Chamaecerasus Rehd. 
Section 1. Isoxylosteum Rehd. 
Subsection 1. Spinosae Rehd. 
13570 =1. L. alberti Rgl. se 478 


Section 2. Isika (Adans.) Rehd. 


Subsection 1. Purpurascentes Rehd. 


2. L. simulatrix Pojark. ae 480 
3. L. microphylla Willd. oe 482 
4. L. pamirica Pojark. bas 484 
Subsection 2. Caeruleae Rehd. 
5. L. kamtschatica (Sevast.) Pojark. at 487 
6. L. pallasii Ldb. ie 489 
7. L. baltica Pojark. see 491 
8. L. edulis Turcz. a 492 
9. L. turczaninowii Pojark. bee 494 
10. L. altaica Pall. ie 496 
13580 11. L. stenantha Pojark. we 497 
12. L. buschiorum Pojark. aut 499 
13. L. caerulea L. oe 500 
14. L. iliensis Pojark. we 502 


XX 


13590 


13600 


13610 


Subsection 3. Chlamydocarpi (Jaub. et Spach) Rehd. 
15,0: 


16. L. 


iberica M.B. 


Subsection 4. Adenostegiae Pojark. 


tolmatchevii Pojark. 


Subsection 5. Bracteatae Hook. f. et Thoms. 


WW 
— 
aT a oa 


Ww 
Nn 
IES} test dest [ext e 


i) 
WwW 
Tail Soll salll alll alll Salll Gall Gall Gall oll soll Gall se 


. hispida Pall. 

. subhispida Nakai 

. semenovii Rgl. 

. heterotricha Pojark. et Zak. 
. praeflorens Batal. 

. altmanni Rgl. et Schmalh. 

. tianshanica Pojark. 


zaravschanica (Rehd.) Pojark. 
bracteolaris Boiss. et Buhse 


. asperifolia Hook. f. et Thoms. 
. olgae Rgl. et Schmalh. 

. humilis Kar. et Kir. 

. cinerea Pojark. 


Subsection 6. Alpigenae Rehd. 


. karelini Bge. 
. glehnii Fr. Schmidt 
. paradoxa Pojark. 


Subsection 7. Rhodanthae Maxim. 


. maximowiczii (Rupr.) Rgl. 

. sachalinensis (Fr. Schmidt) Nakai 
. chamissoi Bge. 

. caucasica Pall. 

. nigra L. 


Section 3. Coeloxylosteum Rehd. 


Subsection 1. Tataricae Rehd. 


L. tatarica L. 

L. micrantha Trautv. 
40. L. 

L 

L 


korolkovii Stapf. 


. lanata Pojark. 
. floribunda Boiss. et Buhse 


XXXi 


503 


505 


509 
511 
513 
514 
515 
S17 
518 
519 
521 
522 
524 
525 
529 


530 
532 
534 


535 
537 
539 
540 
544 


545 
547 
548 
550 
551 


XXXll 


Subsection 2. Ochranthae Rehd. 


43. L. chrysantha Turcz. ad 555 
44. L. globiflora (Rupr.) Dippel te 557 
45. L. xylosteum L. ay 559 
46. L. steveniana Fisch. ne 561 
47. L. ruprechtiana Rgl. Me 563 
48. L. maackii Rgl. me 565 
49. L. nummulariifolia Jaub. et Spach he 566 
Section 4. Nintooa (Sweet) Rehd. 
Subsection 1. Longiflorae Rehd. 
*T.. japonica Thunb. ne 568 
Subgenus 2. Periclymenum Rehd. 
Section 1. Eucaprifolium (Spach) Pojark. 
50. L. caprifolium L. a 570 
13620 51. L. etrusca Santi ane 372 
Tribe 5. Diervilleae C.A.M. 
Genus 1402. Weigela Thunb. 
Section 1. Calysphyrum (Bge.) A. DC. 
1. W. praecox (Lemoine) Bailey ee 575 
Section 2. Synanthera Pojark. 
2. W. suavis (Kom.) Bailey i 581 
xxi Section 3. Calyptrostigma (Trautv. et Mey.) Rehd. 
3. W. middendorffiana (Carr.) C. Koch a6 582 
Family CLIII. Adoxaceae Fritsch 
Genus 1403. Adoxa L. 
1. A. moschatellina L. de 584 


Family CLIV. Valerianaceae DC. 
Tribe 1. Patrinieae Hoeck 
Genus 1404. Patrinia Juss. 
1. P. scabiosifolia Fisch. a 588 


13630 


13640 


13650 


— 


wn 


pede dad 


asl ae} meh as) 


<< 


SSN SNS SN SN SNS SS SS NS SS Se 


. sibirica (L.) Juss. 

. rupestris (Pall.) Dufr. 

. intermedia (Horn.) Roem. et Schult. 
. gibbosa Maxim. 


Tribe 2. Valerianeae (Hoeck) Graebn. 


Genus 1405. Valeriana L. 
Subgenus 1. Valeriana 


Section 1. Tuberosae (Hoeck) Grub. 


. tuberosa L. 
. leucophaea DC. 
. chionophila Pop. et Kult. 


Section 2. Rhizophorae Grub. 


. alliariifolia Adams 
. tiliifolia Troitzky 
. alpestris Stev. 


saxicola C.A.M. 
jJelenevskyi P. Smirn. 
tripteris L. 

dicica L. 


. simplicifolia Kabath 


turczaninovii Grub. 


. capitata Pall. 


sisymbriifolia Vahl 
ficariifolia Boiss. 


. eriophylla (Ldb.) Utk. 


cardamines M.B. 
daghestanica Rupr. 
officinalis L. 

dubia Bge. 


. colchica Utk. 


amurensis P. Smirn. 


. ajanensis (Rgl. et Til.) Kom. 
. petrophila Bge. 

. martjanovii Kryl. 

EY. 


fedtschenkoi Coincy 


XXX1il 


589 
590 
590 
593 


600 
601 
601 


602 
605 
607 
608 
609 
609 
611 
612 
615 
616 
617 
618 
619 
620 
624 
625 
630 
632 
635 
637 
637 
638 
639 


XXii 


XXXIV 


13660 


13670 


P= 


10. 
if. 
jis 
13. 
14. 
15: 


16. 


17. 
18. 


19. 
20. 


SO CLES Me ee 


SSS 55 5 


Sen NN SNS 


Genus 1406. Centranthus Neck. 


Section 1. Macrocentron Lange 


. longiflorus Stev. 
. ruber (L.) DC. 


Section 2. Calcitrapa Lange 


. calcitrapa (L.) Dufr. 


Genus 1407. Valerianella Mill. 


Section 1. Psilocoelae DC. 


. tuberculata Boiss. 
. uncinata (M.B.) Duftr. 
. dactylophylla Boiss. et Hohen. 


turkestanica Rgl. et Schmalh. 


. triplaris Boiss. et Buhse 


diodon Boiss. 


. kulabensis Lipsky. 
. oxyrrhyncha Fisch. et Mey. 


anodon Lincz. 


Section 2. Sclerocarpae Boiss. 


sclerocarpa Fisch. et Mey. 
corniculata C.A.M. 
cymbocarpa C.A.M. 
plagiostephana Fisch. et Mey. 
szovitsiana Fisch. et Mey. _ 
vvedenskyi Lincz. 


Section 3. Cornigerae Soy-Willem. 


echinata (L.) DC. 


Section 4. Locustae DC. 


. locusta (L.) Betcke 
. costata (Stev.) Betcke 


Section 5. Siphonocoelae Soy-Willem. 


. dentata (L.) Poll. 
. muricata (Stev.) W. Baxt. 


640 
641 


641 


647 
648 
651 
652 
652 
653 
654 
655 
656 


656 
657 
658 
659 
660 
661 


662 


663 
664 


667 
668 


XXIil 


13680 


13692 


13694 


SS SS SS 


Genus 1408. Pseudobetckea (Hoeck) Lincz. 
12 


Ss See SS OS 


Section 6. Platycoelae DC. 


. carinata Loisel 
. turgida (Stev.) Betcke 


rimosa Bast. 

amblyotis Fisch. et Mey. 
lipskyi Lincz. 
platycarpa Trautv. 


. pumila (L.) DC. 
. brachystephana (Ten.) Bertol. 
. lasiocarpa (Stev.) Betcke 


pontica Lipsky 


Section 7. Coronatae Boiss. 


. coronata (L.) DC. 

. kotschyi Boiss. 

. dufresnia Bge. 

. leiocarpa (C. Koch) Ktze. 
. vesicaria (L.) Moench 


caucasica (Boiss.) Lincz. 


XXXV 


670 
671 
672 
673 
673 
674 
674 
675 
676 
676 


677 
678 
679 
680 
680 


681 


siete 3D, Celene Roy Wade 


cchinets (1) DOC ie ‘1 6 Oe 


DD “he “ 


Becton 4. Locustse DC) / N 
‘ a : o \ , 
vn he Ye <a } Betke ‘ 0 oan a 


‘ iy 
a 


¥ 


Me 


om 


ot 
i 


{ 


iy fi 


1 


Family CXLIII. BIGNONIACEAE’ PERS. 


Flowers bisexual, irregular. Calyx campanulate, apparently trun- 


cated or 5-toothed (even partite) along margin. Corolla usually some- 
what bilabiate, infundibuliform-campanulate, 5-lobed above. Stamens 
2-4, inserted in lower part of corolla tube; staminodes sometimes (1— 
3). Ovary superior, uni- or bilocular, with numerous ovules; style bi- 
lobed at tip, usually with honey disk at base of ovary. Fruit a capsule, 
sometimes fleshy, nondehiscent. Seeds nonendospermic, almost always 
with winged appendages. Embryo with broad cotyledons, often emar- 
ginate at tip, up to bilobed. Trees, shrubs, sometimes lianas or herbs, 
with opposite or alternate, often compound leaves. Of the more than 
100 genera of this family, the USSR has 4. 


Pe + = 


- 


KEY TO GENERA 


Plant climbing; leaves pinnate, with broad, ovate, serrate-dentate 


EEE OR in ena ee Eee *Campsis Lour. 
Sfemsenect MOL CUMING our asocse ncncmenc arts an nog iano esiutnnoua doisiee suds 2: 
Trees with entire, large leaves ..............::sssceeseesseeees *Catalpa Scop. 
Herbs with, Gissected eave sie secession oad ceeeatho sedan ap ods 3. 
Fruit an elongated capsule, with thin, coriaceous wingless walls; 
seeds winged, wings translucent ................. 1367. Incarvillea Juss. 


Fruit winged capsule, with woody walls; seeds not winged 
ae ge Anas ane ae 1368. Niedzwedzkia B. Fedtsch. 


GENUS *Campsis? Lour. 


Lour. Fl. Cochinch. III (1790) 377 


Flowers large, in terminal, umbelliform racemes. Calyx tubular- 


campanulate, coriaceous, 5-toothed, weekly bilobed. Corolla infundibu- 
2 liform-campanulate, 5-lobed, scarlet or orange. Stamens 4 (didy- 
namous) with curved filaments. Ovary bilocular, surrounded by broad 
disk at base. Fruits— elongated capsule, dehiscing by valves, separated 


'Treatment by I.T. Vassilczenko. 
7From the Greek kampsis—curved (so named because of curved filaments). 


2 


from septum supporting seeds. Seeds numerous, compressed, with two 
translucent, winged appendages. Lianas, climbing with the help of 
suckers (aerial roots), with opposite, odd-pinnate leaves, with dentate 
leaflets. 

Two species are known in this genus, distributed in tropical and 
subtropical Asia and North America. Type of genus: C. radicans (L.) 
Seem. 

Economic importance. Species of this genus are valuable orna- 
mental plants. 


1. C. radicans (L.) Seem. in Journ. Bot. V (1867) 372; Bureau, 
Monogr. Bignon. 145; Schumann in Pflanzenfam. IV, 3b, 230; Vilmor. 
Ill. Blumengart. I, 801; C.K. Schn. Handb. Laubholzk. II, 623; Britton 
and Brown, FI. North. Amer. III, 237; Bailey, Manual of cultiv. Plants. 
903.—Bignonia radicans (L.) Juss. Gen. pl. (1774) 139.—Ie.: Bureau, 
Atlas, pl. 14; Britt. and Brown, I.c. 237. 

Cultivated. Stem climbing, up to 10 m long. Leaves odd-pinnate, 
with up to 7 cm long petiole, with (7)9-11 ovate or oblong-ovate 
leaflets, 4-6(12) cm long, 2—3(7) cm broad; leaflets coarsely serrate- 
dentate to incised along margin, acuminate, dark green above, lighter 
beneath and hairy (at least along midrib). Flowers on terminal branches 
in reduced (umbelliform) racemes. Calyx glandular-pubescent, 5- 
toothed to incised. Corolla usually orange with scarlet limb, 6-8 cm 
long (limb up to 3-4 cm long), 5-lobed above. Ovary with elongated 
style and bilaminate stigma. Capsule oblong-cylindrical, 8—15(20) cm 
long. Keeled along sutures, beaked at tip. Seeds orbicular, with two 
translucent, light brown wings. July to September. 

Cultivated as a prize ornamental plant in the south of the USSR 
(flowers and fruits in the Kiev Region of the European part of the 
USSR). General distribution: North America (southern part). Culti- 
vated in all parts of the world. Described from America. Type in 
London. 

Note. The second species of this genus, C. sinensis (Lam.) Voss. 
in Vilmor. [Ill]. Blumengart. I (1896) 801.—Bignonia sinensis Lam. 
Encycl. I (1783) 423.—B. grandiflora Thunb. Fl. Jap. (1784) 253], is 
also found occasionally in cultivation in the southern part of USSR. 
This species (in contrast with C. radicans) has a shorter but [rather] 
large corolla (up to 5 cm in diameter), leaflets glabrous beneath, calyx 
without glandular hairs, 5-partite, with teeth longer than the tube. This 
plant grows wild in southern China and Japan. 


GENUS 1367. Incarvillea! Juss. 


Juss. Gen. pl. (1789) 138. 

Corolla infundibuliform, large, 5-lobed, slightly bilabiate, pinkish 
purple, pink or white. Calyx campanulate. Stamens 4. Ovary superior, 
bilocular. Fruit—elongated, many-seeded capsule, with coriaceous 
(rarely thicker) valves. Seeds with white, translucent wing. Perennial 
(sometimes annual or biennial), glabrous herbs with opposite or alternate 
pinnatisect (or partite) leaves. 

This genus includes 10-12 species, distributed mostly in China. 2 
species are found in USSR. Type I. sinensis Lam. 

Economic importance. Species of the genus /ncarvillea are valu- 
able ornamental plants. 


1. | Leaves (lower and middle) opposite, pinnatisect ..... 1. I. olgae Rel. 
+ Leaves alternate, bipinnatisect ................ cee 2. I. sinensis Lam. 


1. I. olgae Rgl. in Gartenflora (1880) 3; Descr. pl. nov. rar. 62; 
Fedch. Rastit. Turkest. 700; O. and B. Fedch. Perech. rast. Turkest. V, 
124.—I. koopmanni Lauche, Deutsch. Gart. (1880) 39.—Ic.: 
Gartenflora (1880) tab. 1001 (ic. color.); Lauche, l.c. (ic. color.).— 
Exs.: Gerb. Fl. SSSR (1949) No. 3490; Herb. El. As. Med. (1928) No. 
445. 

Perennial. Stem up to 1-1.5 m tall, sometimes woody at base, 
glabrous, branched above. Leaves opposite, pinnatisect or pinnate, 
uppermost leaves usually entire, oblong-linear; leaf segments (and 
leaflets) oblong-linear or oblong, entire or subdentate, to incised. 
Flowers in lax, large, terminal panicle with opposite, distant branches. 
Pedicels slender, longer than calyx, with small, linear bracteoles at 
base of panicle. Calyx campanulate, glabrous, 3-4 mm long; teeth 
broadly deltoid, 0.6—0.9 mm long. Corolla pinkish purple, rarely white, 
2.5-3 cm long, tubular-cylindrical, broadened above, with 5 orbicular 
lobes (2 of them larger), about 1—1.5(4) cm across. Capsule coriaceous, 
linear, slightly curved, 5—10 cm long, with thick, oblique, intersecting 
veins, acuminate, usually dehiscing by ventral suture (like follicles), 
with thickened septum. Seeds oblong-ovate, compressed, with round, 
white, translucent wing, about 10 mm long. June to October. (Plate I). 

Gravelly valleys, sandy deposits, stony and clayey slopes, at 700— 
800 to 2500 m altitude.—Soviet Central Asia: Pamiro-Alai. Endemic. 
Described from northern spurs of Alai Range. Type in Leningrad. 


‘Named after the French traveler Incarville, who studied the Chinese flora in the 
17th century. 


Note. W. Lauche raised plants in northern Germany from seeds 
obtained from horticulturist Koopmann from Margelan, which he de- 
scribed as a new species, I. koopmanni W. Lauche (l.c.). According to 
Lauche, this species is distinguished by the larger corolla (limb up to 
4 cm across), corolla tube slightly curved at base (this feature is ob- 
served also in J. olgae) and large, bilobed stigma. However, in this 
case one assumes that the larger corolla and stigma resulted from grow- 
ing the plant in soil rich in nutrients and from good care (as mentioned 
by Lauche himself). In view of this, it would be more correct to con- 
sider I. koopmanni as a cultivated form of I. olgae, although such a 
rapid change in the size of the flower under culture conditions in this 
case is a very interesting fact. Also valuable is the information given 
by Lauche about the successful cultivation of J. olgae in northern 
Germany, where this plant withstood wintering very well and flowered 
already in the second year (August-September). In the first year, it 
reached 40-50 cm in height and in the second year, up to 1 m. Further, 
we can add that I.A. Raikova discovered J. olgae in the Pishkharv 
River valley (right tributary of Pyandzh in western Pamir) at an altitude 
of 1700 m, with the flowers having a pink and white corolla. 


2. I. sinensis Lam. Encycl. méth. III (1789) 243; DC. Prodr. IX, 
237. Batalin in Tr. SPb. Bot. sada, XIV, 8, 179; Kom. Fl. Manchzh. 
III, 444; Kom. and Alis. Opred. rast. Dalnevost, kr. II, .934.—I. 
chinensis Poir. in Dict. Sc. Nat. XXIII (1822) 152. 

Perennial. Stem erect, rough, simple or branched, up to 0.5—1 m 
tall. Leaves alternate (upper sometimes opposite), bipinnatisect, with 
narrow, lanceolate-linear or linear segments; segments entire or re- 
motely denticulate. Flowers in apical, lax, leafy racemes or solitary, 
with linear-lanceolate bracteoles at base. Calyx lobes subulate, with 
thickened rounded base, 2-3 times as long as campanulate tube, finely 
glandular-asperous, with reduced (1-1.5 mm long), deltoid-lanceolate 
teeth between long calyx teeth, sometimes bipartite at tip. Corolla 
infundibuliform, 30-50 mm long, 3—4 times as long as calyx, 5-lobed, 
bright purple. Fruit (5)7-10(12) cm long, surrounded at base by 
remnants of calyx, on thickened, short (3-5 mm) stalks, erect, linear- 
lanceolate, usually slightly curved. Seeds ovate, 3-4(6) mm long, with 
white, hyaline wing along margin. Flowering from June. Fruiting from 
- July. 

River valleys, slopes, sometimes old fields, edges, etc., in sands, 
sandy and stony soils.— Soviet Far East: Ussuri. General distribution: 
Mongolia, northern China. Described from environs of Peking. Type 
in Paris. 


=) 


5 


Note. V.L. Komarov and E.N. Alisova (l.c.) report this species 
from the middle reaches of the Sungari River. The Herbarium of the 
Botanical Institute of Akad. Nauk SSSR has specimens of this plant 
collected near the city of Tsitsikar. The discovery of a species of this 
genus (without flowers and fruits) on rocks on the banks of the Suifun 
River within the USSR has been confirmed. But this plant is distin- 
guished from typical biennial J. sinensis by its perennial habit. This 
was the reason given by the above-mentioned authors for treating the 
Suifun Incarvillea as a distinct species (“J. unnamed—lI. sp.”). How- 
ever, since all the remaining features of this Incarvillea sp. are the 
same as in I. sinensis, I consider it best to assign the Incarvillea sp. 
to I. sinensis. More comprehensive material is needed for a final so- 
lution of this problem. 


GENUS 1368. Niedzwedzkia! B. Fedtsch. 


Fedtsch. in Izv. Bot. sada, XV, 3-4 (1915) 1; Rastit: Turkest. (1915) 
701. 

Flowers solitary, large, pedicellate, terminal on stem and branches. 
Calyx broadly campanulate, deeply 5-lobed. Corolla much longer than 
calyx, tubular-infundibuliform, with 5-lobed limb. Stamens 4. Style 
filiform. Capsule hard, with five winged, longitudinal, unequally crenate 
ribs along margin. Seeds ovate, compressed, numerous, with central 
embryo, surrounded by thin layer of nutrient tissue. Semishrub, with 
alternate, dissected leaves. 

One species is known in this genus. 


'This plant has been named in honor of V.E. Niedzwetskii, whose student, Lyutik, 
while participating in his expedition, discovered and collected it. N.A. Bush [in 
Zhurn. Russk. Bot. obsch., III (1918) 153] points out the incorrectness of the form 
of the generic name Niedzwedzkia, which, however, was spelled differently by B.A. 
Fedtschenko himself (in “Rastitel’nost Turkestana” in key on p. 701 as “Niedzwieckya” 
and below on the same page as “Niedzwedzkia”). Bush considers the transcription 
“Niedzwieckia” to be more correct, also pointing out the inappropriate construction of 
the specific name of this plant, which it would have been more correct to name “N. 
heptapotamica.” Furthermore, we may add that the name “N. semiretschenskia” figured 
in the “International Code of Botanical Nomenclature” as an example of violation of 
one of the recommendations (to avoid long words difficult to pronounce while forming 
specific epithets); see also note in V.L. Komarov (Uchenie o vide u rastenii (The 
Species Concept in Plants), 1944). 


f=) 


1. N. semiretschenskia B. Fedtsch. in Izv. Bot. sada, XV, 3-4 
(1915) 399; Rastit. Turkest. 702. —Exs.: Herb. Fl. As. Med. No. 446.— 
Ic.: Fedtsch. in Izv. Bot. sada, XV, 3—4 (color illustration’). 

Perennial. Stems woody at base, ascending, usually several, 20-30 
cm tall, simple or weakly branched in upper part. Leaves numerous, 
alternate, pinnately, or almost palmately dissected into narrow, linear, 
somewhat obtuse lobes. Flowers solitary with leaflike, 3-partite 
bracteoles. Calyx broadly campanulate, dissected up to middle or 
deeper; calyx lobes obovate, mucronate, or ovate, unequal, 3—-5(6) mm 
long. Corolla purplish violet, tubular-infundibuliform, tube 20-30 mm 
long, limb 5-8 mm long, with 5 orbicular lobes. Stamens 4, inserted 
in corolla tube. Ovary elongated, narrowly oblong; style filiform, with 
broadened stigma. Capsule lanceolate, hard, 30-40 mm long, about 15 
mm broad, with four winged, unevenly crenate ribs, bilocular. Seeds 
ovate, flat, covered (especially along margin) with minute papillae, 
rugose-papillulose. May to June. 

River valleys (?) in fringed wormwood semidesert zone (with 
Artemisia maritima s.1.).—Soviet Central Asia: Tien Shan (Chu-Ili 
Mountains). Described from Kildzhan-Saz Ravine (basin of Kopal 
River). Type in Leningrad. 

Note. V.S. Titov, despite careful search, could not find this plant 
in the reported place (“locus classicus’’). In this regard, A.I. Vvedensky 
(Herb. Fl. As. Med.) suggested that the place of initial discovery of N. 
semiretschenskia was incorrectly reported by Niedzwetskii, Titov and 
Ioffe found Niedzwedzkia in Aiderke (“on slopes of hills”). It was also 
collected in the Akrakai massif in 1926 by a collector unknown to me 
(nature of habitat not mentioned). The above-mentioned places in the 
Chu-Ili Mountains represent all the localities of this remarkable plant 
known at present. V.P. Goloskokov does not cite N. semiretschenskia 
B. Fedtsch. in the list of species in his latest checklist of the flora of 
Chu-Ili Mountains (Izv. Akad. Nauk KazSSR, 4 (1949) 65). 

B.A. Fedtschenko (l.c.), while describing this plant, noted that, 
although N. semiretschenskia resembles certain Pedaliaceae in its habit, 
it is nevertheless closer to representatives of the Bignoniaceae (at 
least to Incarvillea olgae from Turkestan). However, in Fedtschenko’s 
opinion, the structure of the seeds and fruit of Niedzwedzkia is very 
different from that which we observe in the Bignoniaceae (particularly 
in I. olgae). In this respect, Niedzwedzkia is rather close to Pedaliaceae. 
Besides, Fedtschenko established the presence of stalked glands in 


Titov (Herb. Fl. As. Med. No. 446) considers that the color of the Niedzwedzkia 
flower is incorrectly shown in the illustration in B.A. -Fedtschenko’s work: it is too 
red, whereas in reality flowers of this plant are purplish violet. 


i 


Niedzwedzkia, similar to-those in representatives of the latter family. 
Thus, this question seemed to be resolving in favor of the inclusion 
of N. semiretschenskia in the family Pedaliaceae. However, A.I. 
Vvedensky (in sched. 1928), having restudied this question, discovered 
that the similarity of Niedzwedzkia to Incarvillea is not limited only to 
the general habit of both species, but extends even further, namely, 
that both genera have stalked glands of identical form. As regards 
seeds, some Chinese species of Incarvillea (e.g. I. compacta Maxim.) 
have seeds similar to those of Niedzwedzkia. Besides, even the fruits 
of I. compacta with rough woody walls are very similar to the fruits 
of N. semiretschenskia, differing perhaps only by the presence of elon- 
gate wings. In view of this, I believe it to be more correct to assign 
Niedzwedzkia to the family Bignoniaceae, where this genus finds a 
place along side Chinese representatives of this family (Incarvillea). 
More detailed justification of this change will be provided later in a 
separate publication. 


GENUS *Catalpa! Scop. 


Scop. Introd. ad Hist. Nat. (1777) 177. 

Inflorescence paniculate or racemose, large. Corolla campanulate 
or broadly tubular, bilabiate, with two upper teeth narrower, three 
lower teeth broad, variously colored (white, pink, yellow, usually with 
stripes and spots of another color), large. Calyx unequally divided, or 
bilabiate, or bidentate. Stamens 4, two of them fertile (with developed 
anthers). Ovary superior; style bilobed at tip, slightly longer than sta- 
mens. Fruit a long, cylindrical capsule, dehiscing by two valves. Seeds 
flat, oblong, numerous, with tuft of white hairs at each end. Trees with 
broad, branched crown. Leaves simple, ovate-cordate or oblong-ovate, 
entire or remotely dentate, sometimes lobed, fetid on crushing in most 
species. Long-petiolate, opposite or whorled. Up to 25 species are 
known in this genus, distributed in America and East Asia, and culti- 
vated also in other parts of the world. Type of genus C. bignonioides 
Walt. 

Economic importance. Species of this genus are excellent orna- 
mental plants (beautiful flowers, fruits and leaves), and their crown 
provides good shade, although in regions of dry rainless summers, for 
example, in Soviet Central Asia, during the summer their large leaves 
collect much dust, which falls down at the slightest movement of 


'Catalpa—local (North American) Indian name for C. bignonioides (“nomen 
vernaculum’). 


8 


the tree and flows in muddy streams during the first autumn rains. The 
following species, of which C. bignonioides is more common, are cul- 
tivated in southern regions of USSR. The wood of Catalpa is distin- 
guished by its hardness and’ durability. Some of the species have 
medicinal importance. 


1. Base color of corolla yellow; corolla not exceeding 3 cm in length; 
leaves often 3-5 lobed .............ceeescesseeeeeeeee 3. C. ovata G. Don. 
+ Base color of corolla white; corolla larger (not less than 4—5 cm 
long); leaves*always entireties tics acmterersseereste se eceee teceene eee 2 


2. Leaves (abruptly) short-acuminate, fetid .... 1. C. bignonioides Walt. 
+ Leaves bape tapering, with long cusp at tip, odorless 
, SE CIR SR sssseesseesesnseeserenennesses Qe C. speciosa Ward. 


1. C. bignonioides Walt. Fl. Carol. (1788) 64; C.K. Schn. Handb. 
Laubholzk. II, 623; Schumann in Pflanzenfam. IV, 3b, 235; Bailey, 
Manual of cultiv. Plants (1949) 906; Paclt. Synopsis of the Gen. Cat- 
alpa in Condollea, 13, 257.—C. cordifolia Jaume in Duham. Arb. ed. 
nov. II (1835) 13.—C. syringifolia Sims. Bot. Mag. XXVII (1880) 
tab. 1094.—C. catalpa (L.) Karst. Deutsch. Fl. (1880-1883) 927; 
Britton and Brown. Fl. North. Amer. III, 238.—Bignonia catalpa L. 
Sp. pl. (1753) 622.—Iec.: C.K. Schn. l.c. f. 403; Britton and Brown, 
kem238: 

Cultivated. Tree 6-8(15) m tall, with widely spreading crown, 
providing ample shade, somewhat rounded in outline. Leaves opposite 
or whorled, 10-20 cm in diameter, broadly ovate, cordate at base, 
abrupt and short-acuminate at tip, sometimes with 1-2 small, acute 
lobes on sides, densely pubescent beneath (hairs multicellular, appear- 
ing nodulose), glabrous above, leaves fetid when crushed, petioles 
thickened, long. Inflorescence a pyramidal panicle up to 20-25 cm 
long; flowers about 4 cm across; corolla white, with two yellow stripes 
inside, densely covered with brownish purple spots. Capsule long and 
slender, cylindrical, pendulous, 15(20)—35(40) cm long, 7-9 mm broad, 
dehiscing along two smooth, dark (blackish) valves. Seeds oblong- 
linear, narrow (3—4 mm broad), concavo-convex, light grayish brown, 
with tufts of white hairs at both ends. June to July. 

Cultivated as ornamental plant in south of USSR. Northern bound- 
ary of cultivation (in open ground) lies along the line Alma-Ata— 
Dzhambul— Kzyl-Orda— Stalingrad— Dnepropetrovsk—Kiev. Pres- 
ently it is being introduced farther north of this line. General 
distribution: North America (southern regions), widely cultivated in 
other parts of the world. Described from Japan. Type in London. 


Plate I. 
Incarvillea olgae Rgl. Lower and upper parts of plant, back and front sides of 


7 


fruit, seed. 


10 


2. C. speciosa Ward. ex Engelm. in Coult. Bot. Gaz. V (1880) 1; 
C.K. Schn. Handb. Laubholzk. I, 625; Britton and Brown. Fl. North. 
Am. III, 238; Bailey, The Stand. Cyclop. Horticult. II (1922) 684; 
Manual of cultiv. Plants, 906; Paclt, Synopsis of the Gen. Catalpa in 
Candollea, 13, 267.—C. bignonioides var. speciosa Ward ex. Barney 
in Gard. Monthly, XX (1878) 312.—Ie.: C.K. Schn. l.c. f. 403. 

Cultivated. Tree 20-30 m tall (shorter in USSR), with pyramidal 
crown. Leaves opposite, ovate or oblong-ovate, 15—30 cm long, acumi- 
nate, mildly cordate at base, densely pilose beneath, odorless. Inflo- 
rescence up to 15 cm long panicle, comparatively few-flowered. Corolla 
whitish-yellow, with obscure spots, about 5 cm across. Capsule 20— 
30(40) cm long, about 1-5 cm broad with thick walls. Seeds oblong, 
broader than in preceding species (up to 5-8 mm broad). June to July. 

Cultivated as ornamental plant in southern USSR. General distri- 
bution: North America (southern part). Introduced in other countries. 
Described from North America. 


3. C. ovata G. Don, Gen. Hist. IV (1837) 230; C.K. Schn. Handb. 
Laubholzk. II, 625; Bailey, Manual of cultiv. Plants, 966; Paclt. Syn- 
opsis of the Gen. Catalpa in Candollea, 13, 252.—C. kaempferi Sieb. 
and Zucc. in Abh. Ak. Muench. IV, 3 (1846) 142; Schumann in 
Pflanzenfam. IV, 3b, 235; C. Henry Dode in Bull. Soc. Dendrol. France, 
6, 199.—Ic.: Dode, I.c. 198; C.K. Schn. l.c. f. 403. 

Cultivated. Tree 6—10(15) m tall, with broad, branched crown. 
Leaves broadly ovate, abruptly narrowed above into short mucro, cor- 
date at base, 10-20 cm in diameter, with 3-5 acute lobes, glabrous 
above and beneath, hairy. only along veins and at nodes between mid- 
rib and lateral veins. Inflorescence paniculate, 10-15 cm long, with 
numerous strong-smelling flowers. Corolla yellow, not more than 2.5 
cm across, with orange stripes and dark violet spots within. Capsule 
linear-cylindrical, 20—30(40) cm long, about 5—7 mm broad (narrower 
than in other species of the genus), with thin walls. Seeds oblong- 
linear, grayish brown, concavo-convex, with tufts of white hairs at 
both ends, smaller than in cultivated species in our country (about 5 
mm long, excluding length of hairs). April to June (July). 

Cultivated as ornamental plant in southern USSR; it is believed to 
be the most frost resistant species as compared with other species of 
the genus. General distribution: China. Cultivated in other countries. 
Described from Japan and China. Type in London. 

Note. Apart from the species listed above, several hybrids have 
been reported in cultivation (for example, C. bignonioides x C. ovata 
and others). 


11 


Family CXLIV. PEDALIACEAE! LINDL. 


Flowers bisexual, irregular. Calyx 5-partite. Corolla 5-lobed. Sta- 
mens 4(5), didynamous, or two developed and two reduced. Ovary 
superior, consisting of 2—4 carpels, sometimes with false septa, very 
rarely inferior (genus Trapella). Style long; stigmas same in number 
as carpels; ovules one or several in each locule, inserted toward central 
placenta. Fruit a dehiscent or non-dehiscent capsule, nutlet-shaped, 
often with appendages (prickles, aristae, etc.). Seeds with thin layer of 
endosperm and erect embryo. Annual or perennial herbs or shrubs, 
rarely aquatic plants (genus Trapella). Leaves simple, usually dentate 
or sometimes sinuate (even partite and dissected), rarely entire, opposite © 
or alternate. Plant glabrous or pubescent with glandular hairs, muci- 


laginous. 


This family includes 16 genera and 120 species, distributed in 
tropical, subtropical and more rarely in temperate regions. 

Economic importance. Of the representatives of this family, the 
most important is sesame (Sesamum indicum L.), cultivated for high 
quality oil, used in food. 


1. Terrestrial plant, cultivated, with erect stem; fruit a dehiscent cap- 
| Sp et fae etre te ea *Sesamum L. 
+ Aquatic plant, growing wild, with creeping or floating stem; fruit 
indehiscent, with 3—5 long, awnlike appendages ....................... 
nd, ae an aa Eh ay a ea ne 1369. Trapella Oliv. 


GENUS *Sesamum? L. 


L. Sp. pl. (1753) 634. 

Flowers usually pentamerous, axillary. Calyx 5-partite. Corolla 
campanulate, almost bilabiate. Stamens 4(5—10) (one or two of them 
usually underdeveloped and reduced to staminodes). Pedicels short, 
with two bracteoles and spherical gland at base. Ovary superior, 2-4- 
locular (sometimes locules parted into longitudinal halves by false 
septa), with central column and seeds adhering to it; style bilobed. 
Seeds numerous in each locule, in single row. Fruit an elongated, 
dehiscent, compressed, 4-angled capsule. Herbs (annuals or perennials) 


'Treatment by I.T. Vassilczchenko. 
"From Greek sezamon — ancient Greek name of sesame. 


12 


with alternate or opposite, entire or parted (up to dissected) leaves. 
Type of genus: S. indicum L. 

Economic importance. This genus includes the important culti- 
vated plant S. indicum L., yielding high-grade edible oil from seeds. 


1. S. orientale L. Sp. pl. (1763) 634.—S. indicum L. 1.c.; DC. 
Prodr. IX, 249; Boiss. Fl. or IV, 81; Stapf in Pflanzenfam. IV, 3b, 
263; Zaitsev in Tr. po prikl. bot. gen. i sel. XIII, 2, 371; Kul’t fl. 
SSSR, VII, 339.—Ic.: Stapf. I.c. & 100; Kul’t fl. SSSR, VII, figs. 
153-161. 

Annual. Stem 75-100 (rarely 250) cm tall, erect, angular in trans- 
verse section (4-angled or 8-angled), sometimes fasciated, sulcate- 
ribbed, usually pubescent (like entire plant), with multicellular capi- 
tate, short and longer hairs or only with short hairs; color of stem 
green, sometimes somewhat bluish (or stem bluish-mottled, against 
green background). Leaves opposite or (in upper part of plant) alter- 
nate, oblong or lanceolate, entire or subdentate, unequally sinuate, up 
to 3-partite or dissected (in middle part of plant, becoming entire 
above, similarly also in lower part of stem), with rather long petioles. 
Flowers 1—3(4—5) in leaf axils; pedicels with spherical, brown, inflated 
gland at base, pitted above. Calyx densely hairy, with 5 long, 
lanceolate-linear teeth. Corolla densely hairy outside, whitish pink or 
pink (to almost violet), (15)20—30(40) mm long, early-shedding, almost 
bilabiate; lower lip usually 3- (sometimes 5-) lobed, with elongated 
middle lobe; upper lip shorter, with 2 (or 3) lobes. Ovary consisting 
of 2 (or 3) carpels (ssp. bicarpellatum Hilt., which should be considered 
typical), or 4 carpels (ssp. quadricarpellatum Hilt., found in Japan 
and isolated areas of South Asia). Style long, linear-cylindrical, lobes 
oblong-ovate. Capsule puberulent, appressed to stem, 30-50 mm long, 
dehiscing almost to base, with thickened short stalk; capsule valves 
usually transversely rugose. Seeds flat, ovate-pyriform, variously col- 
ored (black, gray, reddish brown, yellow, white, with various transi- 
tional shades), 3-3.5 mm long, 1.7—2 mm broad. June to July. 

Cultivated in south of USSR. General distribution: Cultivated in 
countries of southern latitudes (in north, cultivation of sesame reaches 
to 45—46° N. lat.). Grows wild in southern and southwestern Africa. 
Described from India. Type in London. 

Note. Linnaeus (l.c.) referred plants with entire leaves to S. 
orientale, and those with 3-partite lower leaves to S. indicum. How- 
ever, the shape of the leaves in sesame is extremely variable and does 
not work as a basis for separating S. orientale and S. indicum. 

Economic importance. The seeds of sesame are rich in oil, the 
content of which (per absolute weight of dry matter) varies from 47% 


13 


to 62%. Sesame oil is one of a number of valuable food products. 
Besides, sesame seeds are used for preparing confectionery (halva in 
refined oil, etc.). 


GENUS 1369. Trapella! Oliv. 


Oliv. in Ann. of Bot. II (1888) 79. 

Flowers of two types: closed (cleistogamous) and open 
(chasmogamous), solitary, axillary, on long pedicels. Calyx adnate to 
ovary, 5-partite above and with three appendages (in shape of 
cornicles). Corolla infundibuliform, 5-lobed: lobes orbicular, two of 
them connate. Stamens 4 (two of them sterile, reduced staminodes). 
Ovary inferior, bilocular (one locule very small, sterile), style 
elongated, stigma with two short, horizontal appendages at base. Fruit 
cylindrical, oblong- or linear-cuneate, one-seeded, indehiscent, 
terminating into 3-5 aristae, curved at tip. Seeds linear-quadrangular, 
exalbuminous, embryo with narrow, oblong-linear cotyledons. Perennial 
herbs, inhabiting shallow reservoirs, with stems creeping, rooting in 
silt or floating, leaves opposite, dimorphic, petiolate: lower leaves 
elongated oblong, upper broad. Monotypic genus; the sole 
representative (7. sinensis Oliv.) is found in the Far East. 

Note. The slight similarity of the fruits (and in part the upper 
leaves) of this remarkable plant with the fruits (and leaves) of the 
water chestnut (Trapa L.) served as the basis for Lévéille (1899) to 
include Trapella in the genus Trapa (under the name Trapa antennifer 
Lev.). Léveillé, obviously, did not know about the earlier detailed 
study by Olivier (l.c.), who assigned this plant to the family 
Pedaliaceae. However, even within the latter family, the genus Trapella 
occupies a very special position. Stapf (1895) assigns this genus to 
the separate subfamily Trapelleae, distinguished from the rest of the 
Pedaliaceae by an inferior ovary and several other features, and also 
the general set of characters connected with the aquatic habit. It would 
be more correct, in our opinion, to consider this subfamily as an inde- 
pendent family. 


1. T. sinensis Oliv. in Ann. of Bot. II (1888) 79; Forb. and Hemsl. 
in Journ. Linn. Soc. XXVI, 236; Stapf in Pflanzenfam. IV, 3b, 265; 
Kom. and Alis. Opred. rast. Dal’nevost kr. II, 934; Gltick in Bot. 
Jahrbiich. Bd. 70, 149; Kom. Fl. Man’chzh. III, 445.—T. antennifer 
(Lév.) Gliick, l.c.— Trapa antennifer Lév. in Bull. Acad. Intern. Géogr. 
Bot. (1899) 229.—Ic.: Oliv. l.c. pl. 1595 and in Hook. Ic. pl. 1595. 


'Diminutive of Trapa. 


Plate II. 

15 Trapella sinensis Oliv.: a) general appearance of plant, b) ovary during flowering 
stage, c) anther, d) stigma, e) mature fruit, f) fruit in longitudinal section, g) 
embryo, h) upper part of plant from Primor’e with fruits, i) lower 

part of plant, k) fruit of plant from Primor’e. 


15 


Perennial. Stems prostrate, creeping, rooting at nodes in silt (f. 
terrestris Gliick) or floating (f. natans Gliick), simple or somewhat 
branched, from 20 to 50 (100 and more) cm long. Lower leaves nar- 
rowly oblong, wavy (sinuate), dentate-crenate; upper leaves orbicular- 
deltoid or ovate-cordate, sinuately crenate-dentate. Leaves petiolate, 
with three main branched veins diverging from tip of petiole, glabrous 
or hairy beneath along veins. Only cleistogamous, small flowers known 
in f. terrestris; in f. natans, flowers small, cleistogamous, as well as 
larger (from 15-20 to 25-27 mm long), open (chasmogamous). Calyx 
cylindrical-infundibuliform, 5-partite. Corolla tubular, limb 5-lobed, 
two lobes usually connate, three free. Corolla yellow or pale yellow 
outside, lighter above inside (with sky blue tinge). Fruit (16)18—25 
(27) mm long (including stalk), (3)4-7 mm broad, with 5, up to 30— 
50(70) mm long, slender aristae, often uncinate or circinate at tip (two 
of them sometimes short, prickly, like subulate processes), or 3-4 
aristae; ribs passing over surface of fruit (their number corresponding 
with number of aristae, i.e. usually 5); sometimes fruit with longitu- 
dinal narrow wings, or smooth (or almost so). Flowering from May to 
June. Fruiting from July to September. (Plate II). 

In small lakes, ponds and pools, along river banks.—Soviet Far 
East: Ussuri (northward, to Khabarovsk). General distribution: Japan- 
China, Described from Ishang (northern China). Type in London. 

Note. Léveillé (l.c.), and later Gliick (l.c.), have described in more 
detail the species T. antennifer, distinguished from T. sinensis by the 
presence of five aristae at the tip of the fruit and also by the slightly 
larger size of the fruit (and seeds) and more distinct wings of the fruit. 
According to Gliick, this species occurs in Japan, while T. sinensis is 
widespread in China and rare in Japan. In this regard, the following 
should be mentioned. In Primorye, plants are observed with fruits 
having both three and five aristae (cf. V.L. Komarov, l.c.), and 
sometimes with four or five aristae, two of which are rudimentary, 
being represented only by subulate appendages. The other features 
distinguishing T. antennifer from T. sinensis, which are mentioned by 
Gliick, are also unconvincing. Moreover, it is impossible to demarcate 
these species geographically: both “species” were observed in the same- 
bodies of water. 

Thus, it should be acknowledged that there is not a sufficient 
basis at present to recognize the species T. antennifer. Further collection 
of comprehensive material and more observations is necessary. 


18 


Family CXLV. MARTYNIACEAE! LINK 


Calyx membranous, 5-lobed or 5-toothed, sometimes with one 
longitudinal incision in front. Corolla large, pinkish purple, rarely 
white or yellow, symmetrical, thimblelike campanulate, with long or 
short tube, broadened, campanulate at tip; limb somewhat bilabiate, 5- 
lobed. Fertile stamens 4, rarely 2-5, didynamous and one reduced, all 
inserted in broadened part of corolla tube; anthers oblong. Ovary su- 
perior; pistil one, with long style and bilobed stigma. Fruit a longicorn 
capsule with one pericarp. Seeds pendulous, with thick or thin seed 
coat, rugose. 

Annuals with opposite or alternate leaves. 

This family includes three genera with nine species, distributed in 
the tropical and subtropical zones of America. 


GENUS 1370. Proboscidea? Moench 


Moench, Meth. pl. (1794) 466. 

Flowers large, few (3-6), with long pedicels in terminal racemose 
inflorescence. Bracteoles (2-3) linear or oblong. Calyx membranous, 
5-partite or 5-lobed, with longitudinal incision up to base in front. 
Corolla 3.8-6.5 cm long, oblique, broadened almost from base, 
thimblelike campanulate, yellow, pinkish red or purple often with 
yellow stripes. Stamens 4, didynamous, with filiform filaments, fifth 
undeveloped. Pistil one, with oblong-ovate ovary, long style and 
bilobed stigma. Capsule coriaceous, ovate, elongated, rostrate or 
longicorn with incurved beak or horn, dehiscing along with capsule 
on maturity; inner side of capsule valves covered along whole length 
with cristate, hard, oblique processes. Seeds minute, tuberculate-rugose. 
Annual or perennial herbs; leaves opposite or upper ones alternate. 

This genus is represented by six species, distributed in America 
from Arizona and Texas to Peru and from southern Brazil to La Plata. 

P. louisiana (Mill.) Woot. and Standl. in Contribut. from the Un. 
St. Nat. Herb. vol. 19, 602; C. Kott. in Sov. bot. 1, 41; Grossh. Opred. 
rast. Kavk. 321.—Martynia louisiana Mill. Gard. Dict. ed. 8 (1768) 
No. 3.—M. proboscidea Glox. Obs. Bot. (1785) 14; DC. Prodr. IX, 
253.—lIc.: Bot. Mag. XXVI, tab, 1056. 


'Treatment by S.G. Gorshkova 
*From the Greek proboskis — proboscis, according to the shape of the fruit which 
is long bicornuate at the tip. 


17 


Annual. Plant generally up to 20 cm tall, covered with dense, 
white, multicellular, erect, glandular, long, viscid hairs. Root virgate, 
thick, long. Stem somewhat thick, branched from base, branches long. 
Leaves 3-9 cm long, 2.8-9 cm broad, entire, orbicular or cordate, 
obtuse, rugose, entire, densely ciliate, pubescent on both surfaces, 
especially along veins, with 1.5—9 cm long petioles. Flowers large, 3— 
5(6), with strong smell, on slender, pubescent, 2-3 cm long pedicels, 
in racemes. Bracteoles 2, oblong, 1.2 cm long, 3.5 mm broad, 1/2 as 
long as calyx, subobtuse, hairy, entire, densely ciliate. Calyx cam- 
panulate, membranous, glandular-pubescent, ciliate, 2.5 cm long, 1.3 
cm broad, sometimes reddish in upper part; upper lobe deltoid-conical, 
somewhat elongated, 1 cm long, 0.5 cm broad, obtuse, 2 times as long 
as orbicular lateral lobes; lower lobes 0.6 cm long, 0.8 cm broad, also 
orbicular. Corolla 3.8-5(6.5) cm long, 4.54.8 cm across (limb), gla- 
brous outside, glandular-puberulent inside, pale violet, with light red 
and dark violet dots and with 5 orange stripes along lower inner side; 
tube campanulate, broadened from base, 2.5(3) cm long, 0.4 cm broad 
in lower part and 2 cm in upper part, with almost bilabiate, 5-lobed 
limb; upper lobes orbicular, 1.2 cm long, 1.3 cm broad, lateral lobes 
1.4 cm long, 1.6—1.8 cm broad, middle lobe broad, 1.2—1.4 cm long, 
2 cm broad. Stamens with oblong anthers and 1—1.5 cm long filaments. 
Pistil with oblong-ovate, 3 mm long, 1.5 mm broad, brown ovary, 2 
mm long style and bilobed, 3 mm long stigma. Capsule woody, brown, 
ovate or oblong, with incurved, 8-10 cm long horn, 4-angled, 5-15 
cm long, 1.4 cm broad, with coarsely pitted surface, with two deep 
grooves in middle, dehiscing along with horn by two valves, with 
developed, oblique, cristate, acute processes on inner side of valves. 
Seeds 9 mm long, 6 mm broad, ovate-rhombic, convex on outer side, 
dark brown with faint longitudinal stripes, tuberculate-rugose. 

American plant, found as weed in USSR. Cultivated in a number 
of botanical gardens.— European USSR: Lower Don. Caucasus: Cau- 
casian foothills, eastern Transcaucasia. Soviet Central Asia: Syr Darya 
(Tashkent), mountainous Turkmenia (Ashkhabad). General distribu- 
tion: North America. Described from Louisiana. Type in London. 


Family CXLVI. OROBANCHACEAE!’ LINDL. 


Flowers bisexual, zygomorphic. Calyx gamosepalous, campanu- 
late, tubular or patelliform, with 2-5 teeth or lobes, or incised up to 


'Treatment by I.V. Novopokrovsky; manuscript revised by N.N. Tzvelev at editors’ 
request. 


20 


21 


18 


base into 2 (very rarely 3) lateral, entire or bidentate segments. Co- 
rolla gamopetalous, tubular or campanulate, with 4—5 lobed, bilabiate, 
rarely almost regular limb; upper entire, sinuate or bilobed, lower 
usually 3-lobed, with two longitudinal folds between lobes inside throat. 
Stamens 4, filaments glabrous or hairy, geniculate under anthers; an- 
thers dorsifixed, usually bilocular, anther halve (thecae) parallel or 
diverging at tip, but pointing downward due to incurvation of fila- 
ments, usually acute or mucronate; locules fertile, or one of them 
sterile and variably modified; pollen grains globose or ellipsoid, with 
three longitudinal grooves. Ovary superior, unilocular,. consisting of 
two, rarely three carpels, with 2-6 placentae; placentation perietal; 
ovules usually numerous, anatropous. Style one, apical, with 2—-4- 
lobed or entire stigma. Fruit capsule, dehiscing by two, rarely three 
valves. Seeds numerous, minute, variable in shape; embryo small, 
submerged in endosperm, and not differentiated into organs, consisting 
of few cells, seed coat usually one layer of polygonal cells with 
reticulate or pitted, thickened wall. 

Perennial or annual herbaceous plants without cholorophyll, hence 
not green, with simple or branched stems, covered with alternate scales. 
Inflorescence terminal, spike or raceme, rarely corymbose or reduced 
to single flower. Flowers in axils of bract scales, sessile or on some- 
what long pedicels; often with two more lateral bracteoles, attached to 
calyx or pedicel. Parasitic on roots of flowering plants. 

This family includes 15 genera distributed mainly in the northern 
hemisphere in regions with a warm, temperate climate. In the USSR, 
it is represented by six genera. 

Identification of the species of this family, which is biologically 
very interesting, is very difficult. When collecting representatives of 
the Orobanchaceae, it is important to identify the “host” plant and 
also to note the color pattern of the corolla (its limb and tube), stigma, 
stem and scales in the living condition, since the color of these parts 
changes drastically on drying. 


KEY TO GENERA 


1. Stem with one, very rarely two large flowers; pedicels long; calyx 


cainpanulate <2 d.c0... Site ee ee a eee 1372. Phelypaea Desf. 
2)... lowersysathenred) imiinilorescen Ge teres. ace sconce cae onc cy neo ccnepeee 2. 
2. Inflorescence corymbose, sometimes subcapitate; stem extremely 
reduced, often: umGereroumn 22. eseee copes neeee cee ons conse cvetea es oceaeesaaeee 3: 


+ Inflorescence spike or raceme; stems emerging above ground ... 4. 


22 


19 


3. Calyx gamosepalous, with long tube, 4—5-toothed; bracteoles present; 
Ovary “SCOMprisingZe ewoNNEanpels Tee ae eee eee 
MEAG. ae: BOR tT eT i FRE 1375. Mannagettaea H. Smith 

+ Calyx with two (rarely three) free, linear lobes, very similar to 
bracteoles; actual bracteoles absent; ovary comprising three carpels 
Sco SRR a SR EL NS 1376. Phacellanthus Sieb. and Zucc. 

4. Stamens markedly exserted beyond throat of corolla; upper corolla 
lip subentire, several (usually 3—4) times exceeding much reduced 
3-lobed lower lip; calyx obliquely patelliform, shortly and unequally 
2-5 toothed. Parasitic in alder QrOVES .........ceceessseeseeeeseeeeseeeeess 
WA AOR BL OR bs 1374. Boschniakia C.A.M. 

+ Stamens included; corolla limb with five subequal lobes or bilabi- 
ate; in latter case lower lip equaling upper, or longer; calyx 


GiffeHeHat GARR A RRA OG ER OS at 2), 
5. Calyx lobes (sometimes only anterior) broad, obtuse; corolla lobes 
subequal), 002 AoRS, ARS 1371. Cistanche Hoffmg. and Link. 
+ Calyx lobes or segments acute at tip; corolla limb somewhat 
bilabiate, with unequal lobes ................ccseee 1373. Orobanche L. 


GENUS 1371. Cistanche! Hoffmg. and Link 


Hoffmg. and Link, FI. Port. I (1809) 319; G. Beck, Orobanchaceae 
in Pflanzenr. IV, 261 (1930) 26.—Phelipaea sect. Cistanche Walp. 
Repert. III (1844-1845) 461; Reuter in DC. Prodr. XI, 11; Ldb. FI. 
Ross. III, 314; Boiss. Fl. or. 1V, 500.—Orobanche subgen. Cistanche 
Schmalh. FI. II, 295. 

Flowers in dense, spikes or racemes, sessile or short-pedicellate, 
with or without two bracteoles on sides of calyx, with only one bract 
scale. Calyx gamosepalous, campanulate, with 5 subequal, obtuse lobes, 
rarely with deep sinus on side of inflorescence axis; posterior lobe in 
form of short acute tooth, or absent (in latter case calyx 4-lobed). 
Corolla tubular-campanulate, broadened toward throat, with somewhat 
curved tube; limb 5-lobed, almost regular; lobes broad, similar or 
almost identical. Stamens included, inserted in lower third of corolla 
tube; anthers hairy, anther lobes (thecae) mucronate at tip, rarely obtuse. 
Ovary surrounded at base by circular nectary; placentae 4 (rarely 6 or 
2). Capsule ovoid, slightly compressed on sides, dehiscing by 2, rarely 
3 valves; style persistent. Perennial herbaceous. plants, with thick, 
simple stems, covered with alternate scales. 


‘From Greek kystos—capsule, and the word ending from Orobanche, owing to 
the close similarity in the structure of the capsule with the latter genus. 


23 


20 


This genus includes about 18 species, distributed in the warm and 


comparatively dry, mainly subtropical regions of the Old World, from 
the Pyrenean Peninsula and North Africa to China and the northern 
part of India. 


L. 
+ 


2: 


AE 


Flowers with 2 bracteoles on sides Of CalyX ...........esceseeeseeeeeee 2. 

Bracteoles absent, rarely 1 underdeveloped bracteole present (Sec. 

Cistanchiella G. Beck) ........ 7. C. ridgewayana Aitch. and Hemsl. 

Calyx with 5 subequal lobes, rounded at tip. (Sec. Eucistanche G. 

Beck) i s.0s5e-s0cavanssnguss .nasbinserstien!, saute media sun babe eu Me co eBneead Dertlec i. sos B. 

Calyx deeply incised at back (on side of inflorescence axis) with 
4-5 lobes; posterior lobe in form of small acute tooth or absent; 

other lobes larger; 2 anterior lobes always obtuse at tip, 2 lateral 
often subacute (Sec. Heterocalyx G. Beck) ..........cceeesssscceeeeesseneee 
oi Sewanee satel wnlRuaontall ok se bieeiis aelna lei cea 6. C. fissa (C.A.M.) G. Beck. 
Anther lobes obtuse at both ends; plant glabrous throughout, rarely 
very sparsely pubescent along margin of bract scales ............... 4. 

Anther lobes distinctly pointed at one end (morphologically, upper 
end, although it is pointed downward due to geniculation of fila- 
ment under anther); plant somewhat whitish lanate, at least along 
margin\.of ‘bract scales arid bracteoles}...422.4. 2.20. ce 6. 
Flowers whitish pink, 3:5—4.5 cm long::.2.i2: 2.528%. 2 ee 
opt sia A oilliene A OEE. SRA Oe Ei 3. C. monogolica G. Beck. 
Flowers yellow, but often bluish or violet in limb .................... 5. 

Bract scales highly elongated and narrow, considerably exceeding 
calyx and usually also corolla. Parasitic on species of genus 
Calligonumel. A. cs3eecaiis cue aia eae 2. C. flava (C.A.M.) Korsh. 
Bract scales ovate- or oblong-lanceolate, subobtuse, usually slightly 
exceeding calyx, but much shorter than corolla. Parasitic on species 
of genus Tamarix L. ............... 1. C. tubulosa (Schenk) R. Wight. 
Bract scales lanate only along margin, sometimes subglabrous ..... 
Sactesahatdenge: aacuamien. Mendaed etl. birmenell a. 5. C. ambigua (Bge.) G. Beck.. 
Bract scales throughout or partly lanate (but not along margin) .... 
sbeties.. daugeaeet.cgeeeeenl..soedereen: haem hs 4. C. salsa (C.A.M.) G. Beck. 


Section 1. EucistaNcuHE G. Beck, Monogr. Orob. (1890) 57 and in 


Pflanzenr. IV, 261 (1930) 27.—Calyx with 5 identical obtuse lobes. 
Bracteoles present. 


1. C. tubulosa (Schenk) R. Wight, Ic. pl. Ind. or. ITV (1850) 4, 


tab. 1420 and 1420 bis; G. Beck in Pflanzenr. IV, 261, 32, f. 6, F.— 
K.—Phelipaea tubulosa Schenk, Pl. sp. Aegypt. Arab. (1840) 23; 
Reuter in DC. Prodr. XI, 12; Boiss. Fl. or. IV, 500.—Ic.: Wight, I.c.; 
G. Beck, ke: 


24 


21 


Perennial. Plant 30-100 cm tall, glabrous. Stem thick, usually 1.2 
cm in thickness in middle, 6 cm at base, covered with lanceolate, 
subobtuse up to 2—3 cm long scales. Inflorescence cylindrical, dense, 
rarely somewhat lax, 20-70 cm long. Bract scales ovate- or oblong- 
lanceolate, subobtuse, usually slightly exceeding calyx, but much 
shorter than corolla. Bracteoles oblong-lanceolate, usually slightly 
shorter than calyx. Flowers sessile or lower short-pedicellate. Calyx 
usually 2% times as short as corolla, 11-15 mm long, 1/5-1/3 parted 
into subobtuse or broadly ovate lobes, lobes membranous along margin. 
Corolla 28-40 mm long, tubular-campanulate, pale or bright yellow, 
often with bluish tinge in limb; lobes subequal, orbicular or broadly 
ovate, obtuse. Filaments somewhat hairy at base, inserted in lower 
part of corolla; anthers densely villous, 3.5-5 mm long, anther lobes 
(thecae) ovate, obtuse at both ends. Stigma thick, slightly bilobed. 
Capsule dehiscing by 2 (very rarely 3) valves. May to June. (Plate III, 
fig. 2). 

Parasitic on species of Tamarix L. Mainly in river and lake valleys 
in sandy and pebbly soil.— Soviet Central Asia: Kara Kum, mountain- 
ous Turkmenia, Syr Darya, Pamiro-Alai. General distribution: North 
Africa, Ethiopia, Near East, Iran, India-Himalayas (northern part), 
Dzhungaria-Kashgaria (southern part), Tibet. Described from Egypt. 
Type in Oldenburg (Germany). 


2. C. flava (C.A:M.) Korsh. in Zap. Akad. Nauk, VIII ser. 4 (1896) 
5; Grossh. Fl. Kavk. IV, 7.—C. tubulosa var. flava G. Beck in 
Pflanzenr. IV, 261 (1930) 33.—Phelipaea flava C.A.M. Verzeichn. 
Pfl. Cauc. (1831) 104; Reuter in DC. Prodr. XI, 12; Ldb. Fl. Ross. III, 
314; Boiss. Fl. or. IV, 500.—P. gigantea Kar. in Bull. Soc. Nat. 
Mosc. (1839) 165, nomen.—Ic.: M.P. Petrov in Tr. po prikl. bot. gen. 
i Sel.aserul; 4, 123,fiow 5: 

Perennial. Plant 0.8—2 m tall, glabrous. Stem up to 3 cm thick in 
middle, covered with linear-lanceolate, fairly long (3—5 cm long) scales. 
Inflorescence cylindrical, dense, 20-60 cm long. Bract scales oblong- 
linear, narrow, much exceeding calyx, usually almost equaling corolla 
or even longer; bracteoles linear-lanceolate, usually equaling calyx. 
Flowers sessile or short-pedicellate. Calyx usually 24% times as short 
as corolla, 14—20 mm long, up to 1/4—1/3 of length parted into obtuse 
lobes; lobes membranous along margin. Corolla 30-45 mm long, tu- 
bular-campanulate, yellow, lobes subequal, orbicular, obtuse. Filaments 
more or less hairy at base; anthers densely hairy, 3.5-5 mm long; 
anther lobes ovate, obtuse at both ends. Capsule dehiscing by two 
valves. April to May. 


25 


22 


Parasitic on species of Calligonum L. On sand dunes —Caucasus: 
eastern Transcaucasia. Soviet Central Asia: Kara Kum, Kyzyl Kum, 
Balkhash Region (southern part), Pamiro-Alai (western part). General 
distribution: Iran. Described from Azerbaidzhan (between Baku and 
Salyans). Type in Leningrad. 


3. C. mongolica G. Beck in Pflanzenr. IV, 261 (1930) 34. 

Perennial. Plant 30-70 cm tall, glabrous. Stem 1.5—2 cm thick in 
middle, up to 8 cm at base, covered with lanceolate, up to 3 cm long, 
scales. Inflorescence cylindrical, dense, 15-25 cm long. Bract scales 
ovate-lanceolate, subacute, exceeding calyx and usually 1/2—2/3 as 
long as corolla. Bracteoles oblong-linear, usually shorter than calyx 
lobes. Calyx 2%-—3 times as short as corolla, 12-15 mm long, up to 
1/4—1/3 length parted into broad, obtuse lobes, membranous along 
margin. Corolla 35-45 mm long, pinkish white, lobes almost identical, 
orbicular or broadly ovate. Filaments somewhat hairy at base, inserted 
in lower 1/3 of corolla; anthers densely hairy, 4-6 mm long, lobes 
ovate, obtuse at both ends; stigma slightly bilobed. Capsule bivalved. 
May. 

Host plant not known.—Soviet Central Asia: Pamiro-Alai (?). 
General distribution: Dzhungaria-Kashgaria. Described from Kashgaria 
(Chinese Turkestan) (Kuen-Lun, Russian Range). Type in Leningrad. 

Note. G. Beck assigns to this species a specimen of Cistanche 
from the Berlin Herbarium, collected by R.Yu. Rozhevits in 
Tadzhikistan (sands of Patta-Kum, No. 229, 10 April 1906). However, 
its duplicate with the same label, deposited in the Herbarium of the 
Botanical Institute in Leningrad, is indistinguishable from C. tubulosa 
(Schenk) R. Wight, and I, consequently, place it there. In fact, 
differences in the color of the corolla between these species cannot be 
correctly determined on the basis of dry material. 


4. C. salsa (C.A.M.) G. Beck in Pflanzenfam. IV, 3b (1895) 129 
and in Pflanzenr. IV, 261, 36; Grossh. Fl. Kavk. IV, 7; Kryl. Fl. Zap. 
Sib. X, 2535.—Phelipaea salsa C.A.M. in Ldb. FI. alt. IT (1830) 461; 
Reut. in DC. Prodr. XI, 12; Ldb. Fl. Ross. II, 314; Boiss. Fl. or. IV, 
501.—P. armena C. Koch in Linnaea, XIII (1843) 291; Reut. I.c. 11; 
Ldb. |.c. 314.— Orobanche salsa Kuntze in Tr. SPb. bot. sada, X (1887) 
225; Schmalh. Fl. I, 295.—Ic.: Ldb. Ic. pl. fl. ross. IV, tab. 376. 

Perennial. Plant 10-40 cm tall, somewhat hairy. Stem thick, 5—20 
mm thick in middle, markedly thickened at base, covered with oblong- 
lanceolate, alternate scales. Inflorescence short-cylindrical or cylin- 
drical, sometimes strongly reduced, 5-25 cm long, 5-8 cm broad, 
dense. Bract scales ovate- or oblong-lanceolate, somewhat lanate 


26 


23 


dorsally and along margin, up to 2—3.5 cm long, somewhat longer 
than (about 1.5 times) calyx. Bracteoles linear-oblong, subobtuse, 
almost as long as calyx, lanate along margin. Flowers sessile or 
subsessile. Calyx 242-3 times as short as corolla, 9-14 mm long, up 
to 1/4—1/3 length parted into 5 semiorbicular or broadly ovate lobes, 
lobes ciliate ar whitish lanate along margin, calyx sometimes lanate 
throughout. Corolla 25—35 mm long, obtusely-campanulate, slightly 
bent forward, with light yellow tube and violet limb, sometimes light 
yellow throughout, sparsely hairy only along folds of lower lip; corolla 
lobes almost identical, orbicular or broadly ovate, slightly crenate and 
obscurely ciliate along margin. Filaments hairy at base, inserted in 
lower 1/3 of corolla; anthers densely hairy, 3—4 mm long, anther lobes 
distinctly pointed at tip; stigma thick, slightly sinuate. Capsule 
dehiscing by two, rarely three valves. April to May. 

Parasitic on species of Anabasis L. (A. salsa Benth., A. cretacea 
Pall. etc.) Atriplex L. (A. cana C.A.M., A. verrucifera M.B.), Kalidium 
caspicum (L.) Ung.-Sternb. In alkaline soils and salt marshes, rarely 
in calcareous soils and limestone.—European USSR: Lower Volga. 
Caucasus: Southern and eastern Transcaucasia. Soviet Central Asia: 
All regions. General distribution: Asia Minor, Armenia-Kurdistan, Iran, 
Dzhungaria-Kashgaria, Tibet, Mongolia (southern part). Described from 
eastern Kazakhstan. Type in Leningrad. 


5. C. ambigua (Bge.) G. Beck in Pflanzenr. IV, 261 (1930) 40; 
Novopokr. in Fl. Turkm. VI, 293.—C. trivalvis Korsh. in Zap. Akad. 
Nauk. VIII ser. 4 (1896) 6; G. Beck, l.c. 37.—C. eremodoxa Bornm. 
in Fedde, Repert. XV (1936) 336.—Phelipaea ambigua Bge. in Mém. 
sav. étrang. Ac. Pétersb. VII (1854) 429; Boiss. Fl. or. IV, 501.—P. 
trivalvis Trautv. in Tr. SPb. bot. sada, V (1877) 467. 

Perennial. Plant 20-60 cm tall, somewhat hairy, sometimes 
subglabrous. Stem usually 20 mm thick in middle, much thickened 
toward base, covered with glabrous or subglabrous scales, scales 
oblong-lanceolate in upper part of stem, denser and ovate-lanceolate 
in lower part. Inflorescence cylindrical or oblong (to ovate), 10-30 
cm long, 5-10 cm broad, dense. Bract scales oblong-lanceolate, 
subacute, whitish lanate or ciliate along margin, glabrous dorsally, 
exceeding calyx, more than 1/2 as long as corolla. Bracteoles oblong, 
almost as long as calyx, ciliate along margin or subglabrous; flowers 
sessile or subsessile. Calyx 5-lobed, 24-3 times as short as corolla 
and incised slightly deeper; posterior lobe sometimes smaller than 
others. Corolla 30-45 mm long, somewhat hairy along folds of lower 
lip and often also along margin of lobes; lobes subequal. Filaments 
glabrous or sparsely hairy at base; anthers densely hairy, 3-4 mm 


24 


long, anther lobes distinctly pointed at tip. Capsule dehiscing by three 
or two valves. April to May. (Plate III, fig.1). 

Parasitic on species of saxaul (Haloxylon Bge.), possibly also on 
other Chenopodiaceae. Mainly in sands.—Caucasus: Southern 
Transcaucasia. Soviet Central Asia: Aralo-Caspian Region (southern 
part), Balkhash Region (southern and eastern parts), Kara Kum, Kyzyl 
Kum, Syr Darya, Amu Darya, mountainous Turkmenia, Pamiro-Alai. 
General distribution: Armenia-Kurdistan, Iran, Dzhungaria-Kashgaria, 
Mongolia, Tibet. Described from Kyzyl Kum desert. Type in Paris (?), 
topotype in Leningrad. 

Note. The distinctions between this species and the preceding 
species are not always clearly expressed. Bunge described it from an 
atypical specimen collected by Lehmann. The specimen has a strongly 
reduced posterior calyx lobe, and because of that G. Beck erroneously 
assigned C. ambigua to the following section. 


Section 2. HETEROCALYXx G. Beck, Monogr. Orob. (1890) 57 and in 
Pflanzenr. IV, 261 (1930) 39.—Sect. Cistanchopsis G. Beck in 
Pflanzenfam. IV, 3b (1895) 129.—Calyx deeply cleft on side of 
inflorescence axis, 4—5-lobed; posterior lobe a small acute tooth or 
entirely absent, others larger; two anterior lobes always obtuse, whereas 
2 lateral lobes often subacute. Bracteoles present. 


6. C. fissa (C.A.M.) G. Beck in Pflanzenfam. IV, 3b (1895) 129 
and in Pflanzenr. IV, 261 (1930) 39, f. 7; Grossh. Fl. Kavk. IV, 8.— 
C. sintenisii G. Beck in Bull. herb. Boiss. 2 sér. IV (1904) 686; 
Novopokr. in Fl. Turkm. VI, 294.—C. fissa var. sintenisii G. Beck in 
Pflanzenr. IV, 261 (1930) 40.—Phelipaea fissa C.A.M. Verzeichn. 
Pfl. Cauc. (1831) 104; Reuter in DC. Prodr. XI, 12; Ldb. Fl. Ross. III, 
314; Boiss. Fl. or. IV, 501.—P. trivalvis var. lanuginosa Lipsky in 
Zap. Kievsk. obsch. estestv. XI, 2 (1891) 16.—Ic.: G. Beck, in 
Pflanzenr. IV, 261. 

Perennial. Plant 15—50 cm tall, somewhat lanate. Stem 5-20 mm 
thick in middle, covered. with ovate-lanceolate scales, scales longer 
and more densely lanate in upper part of stem. Inflorescence oblong- 
ovate or cylindrical, dense, 5-20 cm long, 5-8 cm broad. Bract scales 
ovate- or oblong-lanceolate, usually subobtuse, somewhat lanate dor- 
sally and along margin, 1.5-2 times as long as calyx. Bracteoles ob- 
long-linear, almost as long as calyx, lanate or ciliate along margin; 
flowers sessile or subsessile. Calyx 10-14 mm long, usually 1/3 as 
long as corolla, deeply incised on side of inflorescence axis, 4—5- 
lobed; posterior lobe a small acute tooth or entirely absent, others 
larger, two anterior lobes always obtuse, lateral often subacute and 


25 


27 narrower; lobes whitish lanate along margin. Corolla 25-35 mm long, 
tubular-campanulate, mildly bent forward, with whitish tube and bluish 
or violet limb, usually yellowish throughout when dry, lobes almost 
similar, orbicular or broadly ovate, somewhat whitish lanate along 
margin and often also inside. Filaments hairy to subglabrous at base, 
inserted in lower 1/3 of corolla; anthers densely hairy, 3-4 mm long, 
anther lobes distinctly pointed at tip; stigma thick, subentire. Capsule 
dehiscing by two valves. April to May. (Plate III, fig. 3). 

Parasitic on subshrubs of family Chenopodiaceae (in Turkmenia, 
reported on. Salsola aucheri Bge). In alkaline soils and salt marshes, 
mainly clayey and stony soils.—Caucasus: Eastern and southern 
Transcaucasia. Soviet Central Asia: Amu Darya, mountainous 
Turkmenia. General distribution: Armenia-Kurdistan, Iran. Described 
from Azerbaidzhan. Type in Leningrad. 

Note. C. sintenisii G. Beck, distinguished by the more dense pu- 
bescence of the corolla lobes, hardly deserves to be recognized even 
as a variety. There is no reason to report this species for Turkmenia, 
as was done by I.V. Novopokrovsky in “Flora Turkmenii”, since the 
pubescence of the corolla lobes is equally variable, both in Turkmenia 
and Transcaucasia. 


Section 3. CISTANCHIELLA G. Beck in Pflanzenr. IV, 261 (1930) 
40.—Calyx with 5 almost similar lobes. Bracteoles absent (rarely one 
underdeveloped bracteole present). 


7. C. ridgewayana Aitch. and Hemsl. in Trans. Linn. Soc. 2 ser. 
III (1888) 93, tab. 39; G. Beck in Pflanzenr. IV, 261, 40, f. 8.—Ic.: 
Ait. and Hemsl. l.c.; G. Beck, l.c. 

Perennial. Plant 15-35 cm tall. Stem glabrous, 0.5—1 cm thick in 
middle, clavate at base, densely covered with brownish scales; scales 
glabrous or lanate along margin, oblong-lanceolate, longer in upper 
part, about 1.5 cm long. Inflorescence oblong-ovate or cylindrical, 
very dense, 8—20 cm long, up to 5 cm broad. Bract scales oblong- or 
ovate-lanceolate, subobtuse, 15-20 mm long, lanate along margin. 
Bracteoles absent (rarely one bracteole present). Calyx 10-12 mm 
long, 1/4—1/3 incised into obtuse, semiorbicular or broadly ovate lobes, 
glabrous or ciliate along margin. Corolla 23-30 mm long, tubular- 
campanulate, with light yellow tube and chestnut (reddish yellow) 
colored limb, orange-brown when dry; corolla glabrous outside, densely 
hairy inside at base of tube; lobes almost similar, orbicular. Filaments 
lanate at base, inserted in lower 1/3 of corolla; anthers densely hairy, 
3—4.5 mm long; anther lobes distinctly pointed at tip; stigma thick, 


28 


26 


semiglobose. Capsule dehiscing by three, rarely two valves. April to 
May. (Plate III, fig. 4). 

Parasitic on species of Tamarix L. (?).— Soviet Central Asia: Kara 
Kum (Akar-Cheshme), Pamiro-Alai (Kelif and Angor). General distri- 
bution: Iran-Afghanistan. Described from Afghanistan. Type in London. 


GENUS 1372. Phelypaea! Desf. 


Desf. Fl. atlant. II (1800) tab. 60.—Anoplanthus sect. Anblatnum Endl. 
Gen. (1839) 727. 

Flowers solitary (very rarely 2), large, ebracteolate. Calyx 
gamosepalous, campanulate, bilabiate or almost regular, with 5 some- 
what connate lobes, glandular-pubescent. Corolla tube bent forward 
and inflated at base, limb broad, 5-lobed, lobes almost identical, broadly 
ovate to suborbicular; inner side of lower lip with two hairy folds 
between lobes. Stamens 4, included, inserted slightly below middle of 
corolla tube; anther lobes distinctly pointed at tip. Ovary with nectary 
at base; stigma thick, capitate; placentae 4. Capsule ovate, slightly 
compressed on sides, dehiscing by two valves. Perennial herbaceous 
plants with simple stems, covered with few amplexicaul scales and 
transforming above into single-flowered peduncle. 

This genus includes 4 species, distributed in Crimea, Caucasus 
and Near East (including Iran). 


1. Calyx with very short, glandular pubescence, subregular or bilabi- 
ate, lobes always lanceolate or subacute; pubescence almost indis- 
cernible without magnification ...............cssccssecessssceseeesneeesnsesenees 2: 

+ Calyx with longer pubescence, bilabiate; upper lip with three short, 
obtuse teeth at tip; lower lip with two long lobes; pubescence 
distinctly discernible without magnification ...............::ssssesseeseeees 
luc. Eanes. eon te! wee 3. P. tournefortii Desf. 

2. Corolla orange yellow, folds of lower lip red; stem and scales light 
orange, brown when dry .............csssccessseeseeeees 2. P. helenae Popl. 

+ Corolla bright red, folds of lower lip velutinous black; stem and 
scales reddish, reddish brown when dry ...........::csssssstescssessesessesseeeneass 
ae RBA Es bs BREE AE a a, a 1. P. coccinea (M.B.) Poir. 


1. P. coccinea (M.B.) Poir. Encyl. méth. V (1804) 268; Fedch. 


and Fler. Fl. (1910) 887; G. Beck in Pflanzenr. IV, 261, 42, f. 9 (A- 
B); Grossh. Fl. Kavk. IV, 8 and Opred. rast. Kavk. 352.— 


‘Named after French patrons A. and H. Phelipeaux. 


29 


27 


P. biebersteinii Fisch. in herb.—P. foliata Lamb. in Trans. Linn. Soc. 
X (1811) 260, tab. 7; Stapf in Kew Bull. (1915) 292.—P. bayerni 
Novopokr. in Grossh. Opred. rast. Kavk. (1949) 730.—Orobanche 
coccinea M.B. Tabl. prov. Casp. (1798) 58 and FI. taur.-cauc. II, 84; 
Willd. Sp. pl. Ill, 354.—O. phelipaea M.B. Beschreib. Land. Cauc. 
(1800) 179 and Fl. Cauc. III, 179.—Anoplanthus coccineus Walp. 
Repert. III (1844-1845) 481, p.p.; Boiss. Fl. or. IV, 494; Schmalh. FI. 
II, 293.—A. biebersteinii Reuter in DC. Prodr. XI, 42; Ldb. Fl. Ross. 
III, 324.—Ic.: M.B. Cent. pl. rar. ross. II, tab. 56; Lamb. l.c.; Stapf, 
l.c. 294; G. Beck, l.c. 

Perennial. Plant up to 30(50) cm tall, reddish or reddish brown, 
covered with very short, dark, glandular, hairs. Stem terete, fistular, 
3-6 mm thick, with few-(usually up to 6), ovate, amplexicaul scales; 
stem transformed above into single-flowered peduncle (very rarely 
with two flowers). Calyx pubescent with very short, dark, glandular 
hairs (almost indiscernible without magnification); lobes lanceolate, 
subacute, subequal; three posterior lobes connate and shorter. Corolla 
(1.8)2.5—4.5(5) cm long, bright red, with two velutinous black folds 
covered with dense, dark hairs. Anthers glabrous or sparsely pubes- 
cent. May to July. 

Parasitic on species of Psephellus Cass. On stony and rubbly 
slopes.— European USSR: Crimea. Caucasus: All regions. General 
distribution: Near East, Armenia-Kurdistan, Iran. Described from 
Caucasus. Type in Leningrad. 

Note. Cases of partial fusion of the three posterior calyx lobes 
occasionally occur throughout the geographical range of P. coccinea. 
Apparently, P. bayernii Novopokr. was described on the basis of such 
specimens from southern Transcaucasia. In this case, however, the 
possibility of their origin as a result of hybridization between P. 
coccinea and P. tournefortii cannot be excluded. 


2. P. helenae Popl. in Dokl. Akad. Nauk SSSR, (1928) 385.—P. 
coccinea f. aurantiaca G. Beck in Pflanzenr. IV, 261 (1930) 43.—Ic.: 
Poplavskaya in Tr. po izuch. Krymsk. zapovedn. (1931) 88. 

Perennial. Plant up to 30 cm tall, light orange, brown when dry, 
covered with very short, light, glandular hairs. Stem terete, fistular, 
3—6 mm thick, with few ovate, amplexicaul, somewhat distant scales, 
terminating into single-flowered peduncle. Calyx pubescent with very 
short, light, glandular hairs (almost indiscernible without magnifica- 
tion), subregular or bilabiate; three posterior lobes connate and shorter, 
but always subacute. Corolla 25-45 mm long, orange yellow, lighter 
outside, with two reddish folds inside throat at base of lower lip, 


30 


28 


covered with light hairs. Stamens inserted slightly below middle of 
corolla tube, anthers glabrous or subglabrous. May to June. 

Parasitic on Psephellus declinatus (M.B.) C. Koch. On stony 
slopes. — European USSR: Crimea. Endemic. Described from Crimean 
Reserve. Type, apparently, lost; cotype in Leningrad. 

Note. P. helenae Popl. probably is not a separate species, but a 
case of polychromism (different coloration of flowers, in this case 
even of the whole plant, in individual specimens of the same species) 
of P. coccinea (M.B.) Poir. Both yellow and red forms occur together 
on stony mountain slopes of the Crimean Reserve on the same host 
plant and, consequently, are not isolated either ecologically or physi- 
ologically. However, P. helenae, apparently does not occur in the 
Caucasus (the record from Mtskhet—appeared to be an error). 


3. P. tournefortii Desf. in Ann. Mus. Paris, X (1807) 298, tab. 
21; G. Beck in Pflanzenr. IV, 261 (1930) 44, f. 9 (C—G); Grossh. FI. 
Kavk. IV, 8.—Lathraea phelypaea var. B. L. Sp. pl. (1753) 606.— 
Anoplanthus tournefortii Walp. Repert. IIT (1844-1845) 481, p.p.; 
Reuter in DC. Prodr. XI, 42.—A. coccineus var. peduncularis Boiss. 
Fl. or. IV (1879) 494.—Ic.: Desf. l.c.; Beck, l.c. 

Perennial. Plant up to 30 cm tall, reddish or reddish brown, glan- 
dular-Pubescent. Stem terete, fistular, 3-5 mm thick, with few (usually 
two), oblong-ovate, amplexicaul scales only at base, terminating into 
single-flowered peduncle. Calyx puberulent with light, glandular hairs 
discernible with naked eye, bilabiate, upper lip with three short, obtuse 
teeth, lower lip with two long, lanceolate lobes. Corolla 30-50 mm 
long, bright red, lighter and puberulent outside, with two velutinous 
black folds, covered with dense, dark hairs in throat at base of lower 
lip. Stamens inserted near middle of corolla tube; anthers somewhat 
hairy. May to June (Plate III, fig. 5). 

Parasitic on Pyrethrum myriophyllum (Willd.) C.A.M.; reported 
also on species of Anthemis L. On stony slopes.— Caucasus: Southern 
Transcaucasia. General distribution: Armenia-Kurdistan. Described 
from Turkey. Type in Paris. 


GENUS 1373. Orobanche! L. 


L. Sp. pl. (1753) 633, p.p.; G. Beck, Monogr. Orob. (1890) 73 and in 
Pflanzenr. IV, 161 (1930) 44.—Phelipaea auct. p.p. non Desf. 


‘Name used by Dioscorides for one of the parasites on leguminous plants; from 
the Greek: orobos—pea (and, generally, leguminous plants), and anchein—to choke, 
stifle. 


33 


29 


Flowers in spikes or racemes, sessile or short-pedicellate, rarely 
(sec. Gymnocaulis Nutt.) long-pedicellate, with or without two 
bracteoles on sides of calyx, with only one bract scale. Calyx 
gamosepalous, campanulate, with 4—5 acute teeth, often deeply parted 
on side of inflorescence axis or divided up to base along median plane 
into two lateral, entire or bilabiate segments. Corolla tubular, 
campanulate or infundibuliform, somewhat broadened toward throat, 
bilabiate; upper lip bilobed or entire, sometimes galeate; lower lip 3- 
lobed, with two folds between lobes. Stamens included; anthers gla- 
brous or hairy, anther lobes distinctly pointed at tip. Ovary ovate or 
cylindrical; placentae usually 4. Capsule dehiscing by two, rarely three 
valves. Seeds minute, numerous. Herbaceous plants, with thick, simple 
or branched stems, covered with alternate scales and often tuberous or 
clavate at base. Parasitic on roots of flowering, mainly herbaceous 
plants. 

This genus includes about 150 species, distributed mainly in 
subtropical and temperate regions of the northern hemisphere. 

Note. The biology of Orobanche has not been adequately studied 
so far. According to Koch (L. Koch, Die Entwickelungsgeschichte der 
Orobanchen, 1887) and other authors, the species of Orobanche para- 
sitizing perennial plants usually behave as perennials, producing new 
shoots in successive years, provided the host plant does not perish due 
to exhaustion or other causes. According to Koch, these species can 
propagate also vegetatively, forming adventitious buds on secondary 
suckers (haustoria), which later lose the connection with the mother 
plant. Species parasitizing annual plants perish along with them and 
are annuals, although some of them can parasitize perennials also. 
There are also biennial species which develop, in the first year, a 
tuber-like growth on the root of the host plant with primary and 
secondary suckers (haustoria), and a stem with an inflorescence in the 
second year. Many species, apparently, can behave as perennials under 
certain conditions and as biennials or even annuals under other condi- 
tions. Therefore, published reports in this respect are often contradic- 
tory, inaccurate and need confirmation. The taxonomy of Orobanche 
is inadequate at present in many respects. Extensive research in this 
field has been done by the monographer of the genus, G. Beck von 
Mannagetta (1890 and 1930), whose classification is mainly followed 
in the present treatment. However, the material from the territory of 
the USSR available to G. Beck was inadequate; many species, therefore, 
were not known to him. These species are most abundant in the 
Caucasus and Soviet Central Asia, where the total number of species 
undoubtedly exceeds the number already known. When collecting 


30 


SI 


rai SS 
S 


6 
og sa 
P Say oS 
EAE I 2 
= roy iG 
Ore 
———=_ —— = 


Ne 
AX. SS 


Plate III. 


31 1. Cistanche ambigua (Bge.) G. Beck, general appearance of plant, flower, 

anther. —2. C. tubulosa (Schenk) R. Wight, anther.—3. C. fissa (C.A.M.) G. 

Beck, flower.—4. C. ridgewayana Aitch. and Hemsl., flower. —5. Phelypaea 
tournefortii Desf. flower. 


34 


31 


Orobanche, it is important to identify the host plant, as well as the 
color of the corolla in the living condition. 

Economic importance. There are species among Orobanche which 
are harmful to agricultural crops, drastically reducing their yield. They 
are: branched broomrape—O. ramosa L., cabbage broomrape—O. 
brassicae Novopokr., Mutel’s broomrape—O. mutelii F. Schultz and 
Egyptian broomrape—O. aegyptiaca Pers., parasitic on cabbage, to- 
mato, melon, watermelon, cucumber, hemp, tobacco and many other 
crops; sunflower broomrape—O. cumana Wallr., parasitic mainly on 
sunflower; alfalfa broomrape—O. lutea Baumg., parasitic on alfalfa 
and clover. Basic measures in dealing with them are: 1) better 
agrotechniques, 2) regular crop rotation, 3) cultivation of broomrape- 
resistant varieties, 4) weeding of broomrape before the formation of 
seeds and the compulsory burning or deep burial of its inflorescences, 
5) so-called “provocative” crops, i.e., sowing in fields affected by 
broomrape such crops as are resistant to it (for example, maize, 
soybean, flax). The last measure helps in markedly reducing seed 
bank of broomrape in the soil. Of late, various biological methods of 
controlling broomrape are under trial (the fungus Fusarium orobanches, 
the fly Phytomyza orobanchia, and others). For details on the control 
of broomrape, see the brochure by I.G. Beilin [Zarazikhi 1 bor'ba s 
nimi (Broomrapes and their control), (1947)]. See notes under 
individual species. 


1.  Bracteoles present; calyx gamosepalous, campanulate, 4—5-toothed; 
stem simple or branched (Sect. Trionychon Wallr.) ................... 2. 
+  Bracteoles absent; calyx divided up to base into two lateral, entire 
or bidentate segments, rarely gamosepalous, campanulate; stem 


always simple (Sect. Osproleon Wallr.) ...............:cssscsssessseeseees 33. 
2. Pubescence of inflorescence, at least along back of bract scales, 
lanate on account of arachnoid-villous hairs; stem simple ......... 3. 
+  Pubescence of entire plant short, glandular, often indiscernible 
Without magnification Vak1.. aeRO weed 6. 


3.  Anthers glabrous, rarely with isolated hairs at base ................:ssss008 
AS RAL Soir cee sei S208) telson ts, Ces ols acer Be 3. O. caesia Rchb. 
+ Anthers densely hairy at base, and often also along sutures of 
theca 2, 2. A oeccessnasessnses bth ariel ae ah) ieee, es acne 4. 
4. Corolla pale sky-blue, usually light yellow when dry, 18-23 mm 
long; inflorescence cylindrical, dense and many-flowered (Kopet- 
Dag) ee BIE, sen deatesncicnnneesties 11. O. borissovae Novopokr. 
+ Corolla violet or blue, usually partly retaining color even when 
dry, (20)22—28(30) mm long (Transcaucasia) ...............:.ssceeee 5. 


35 


32 


Ot Sea 


12: 


13% 


14. 


Inflorescence dense, short cylindrical, often subcapitate; calyx teeth 
slender, usually as long asytubet¢... co). in: neces ee ek eee 
Pein, Sear ser tts, 25552 OTR. 4. O. pulchella (C.A.M.) Novopokr. 
Inflorescence usually lax, cylindrical; calyx teeth thicker, usually 
slightly longer than tube .................:ccc08 5. O. bungeana G. Beck. 
Folds of lower corolla lip extending between its lobes in small 
teeth, lower lip as a result appearing 5-lobed; corolla 16-20 mm 
long, with broadly ovate, subacute lobes; inflorescence dense, 
cylindrical (Transcaucasia) ..... 14. QO. septemloba (G. Beck) Tzvel. 
Lower corolla lip without teeth formed by folds, 3-lobed ........ te 
Folds of lower corolla lip entirely glabrous; corolla 18-20 mm 
long; calyx teeth much shorter than tube, deltoid at base, short- 
subulate pointed; inflorescence dense, many-flowered ............ 
SEU Aceon Skee MARAE RIES. iG Na eh 12. O. kelleri Novopokr. 
Folds of lower lip somewhat hairy 0.0.0.0... ces tsscssecereeeseeeeeeeeees 8. 
Corolla (10)12-15(17) mm long; stem often branched .............. 9. 
Corolla more than: 17 mm long e2.2on.. e)..0.6 Do fees 14. 
Lobes of lower corolla lip obtuse, orbicular or broadly ovate; corolla 
usuallys10=L5 immvlong o23}.).2e tebas. 22s. See Ses 10. 
Lobes of lower corolla lip subacute, ovate; coiolla usually 15-18 
mm longjese..) .uied..22.)..¢4..cmdowd a See eee 11. 
Stem usually branched; inflorescence lax, with spaced flowers; 
corolladimb pale violet. .....:cic<.-...+-2.s.)cecmens oooh 27. O. ramosa L. 
Stem usually simple, rarely branched; inflorescence dense; corolla 
limbjusually violet:..22¢22e ies. xeln eee 26. O. nana Noé. 
Inflorescence lax, relatively few-flowered; calyx teeth usually shorter 
than tube; lobes of lower corolla lip crenate (Crimea, Transcausasia) 
Stee Se SEES. ssissssseseserssesssessseeeseeeee 22. Q, OXYloba (Reut.) G. Beck. 
Inflorescence many flowered, dense, but lower flowers usually 
distant; calyx teeth usually longer than tube; lobes of lower corolla 
lipsnarrower,; subentire s-.1iv. intakes eee ee eee 12. 
Lower flowers usually distantly spaced at anthesis .................. 13. 
Inflorescence dense throughout, rarely lower flowers slightly distant 
(Transcaucasia) ...........::ccssccesceeees 25. O. dalmatica (G. Beck) Tzvel. 
Pubescence of corolla short outside, but discernible without mag- 
nification; filaments of stamens diffusely hairy at base; stem gray- 
ish, glandular-hairy toward tip ............. 23. O. orientalis G. Beck. 
Pubescence of corolla very short, discernible only through magni- 
fying lens; filaments glabrous; pubescence of stem indiscernible 
without magnification ...............sccsseeeeee 24. O. sogdiana Novopokr. 
Calyx teeth and tube somewhat serrate or dentate along 
margin; corolla 20-25 mm long; inflorescence dense (southern 
Transcaucasia) .....2224 2 2a 15. O. serratocalyx G. Beck. 
Calyx tube'and teeth entire ..222o oon cocs cece ons ence nen ececvcousaeessecevscsenes 15. 


36 


37 


LS: 


18. 


33 


Calyx teeth long, setaceous at tip, 142-24 times as long as tube 


(between lateral teeth) (Transcaucasia) .............ssssseseeeseeeees 16. 
Calyx teeth shorter, equaling or not more than 1% times as long as 
FDS ee, FR i BRR A cccwescsinasstisreenasree BR bossonese 18. 


Corolla 25—30 mm long; inflorescence 1aXx ...........cscseeseesesseeeeeees 
WUMARS ED BAS Misi Mi atekctec 16. O. hirtiflora (Reut.) Tzvel. 
Corolla 18-20 mm long; inflorescence dense ..............:c:s+0 ie 
Stem usually branched in anthesis; calyx divided almost up to base 
into 4 teeth on side of infloresCeNCe ..............cccceseeescesteeseeeeeteeees 
Se RIE PES RAR Laces cases 20. O. hohenackeri (Reut.) Tzvel. 
Stem simple; calyx parted on side of inflorescence axis, with 2—2.5 
mm deep incision not reaching its base and almost always with 
short Sth tooth here .................006 10. O. coelestis Boiss. and Reut. 
Inflorescence lax (but dense at tip even at flowering stage), usually 
longer than rest of stem; stem usually branched, often flexuous, 
with few ovate or ovate-lanceolate Scales ............c:scescseseereereees 19. 
Inflorescence dense or lax, in latter case much shorter than rest of 
stem; stem simple covered with rather numerous lanceolate scales 


SERRE, RUNCORN S PMI CRIN OL SID. Sach vohiel eeasedeatee Roark stent aetak 24. 
Lobes of lower corolla lip ovate, subacute, corolla up to 18 mm, 
farely 20 mini long See EAS Ae eco ene 11'. 
Lobes of lower corolla lip orbicular or broadly ovate, obtuse, if 
lobes subacute, corolla larger ..............cccccessceseesssseseeeeteceeneeeesees 20. 
Corolla 24-35 mm long, limb markedly broadly infundibuliform 
PORTIS, AD oh. sec snoscsewwraetirerestereaeasuancast tees raaaaate heer aR EEC cate 21. 
Corolla up to 22, rarely 24 mm ION ...........cccseeeseeeteeeeeeseeees 22. 


Calyx brownish; plant densely pubescent throughout, pubescence 
discernible without magnification (southern Transcaucasia) .......... 
A SRS EET, SI 16. O. hirtiflora (Reut.) Tzvel. 
Calyx lighter, often whitish, chartaceous; pubescence very short, 
indiscernible without magnification .......... 17. O. aegyptiaca Pers. 
Anthers glabrous, rarely sparsely hairy at base (south of European 
part GPUSSR)".. PE. LA RE 19. O. brassicae Novopokr. 
Anthers distinctly and usually densely hairy at base ............... 23° 
Inflorescence starting almost from soil surface (Pamiro-Alai) ..... 
RO ne a ee ee 21. O. brachypoda Novopokr. 
Part of stem carrying inflorescence longer (Crimea, Caucasus) ..... 
REO oN Pet edve is. BARE LE, ROS 18. O. mutelii F. Schultz. 
Corolla with slightly bluish tinge only in limb, light yellow 
throughout when dry (Soviet Central Asia) ............:eeeeeeeeeees 25. 


‘Referral back to couplet 11 is given, because in the species belonging here, the 


corolla can be up to 20 mm long. 


38 


34 


27. 


31. 


a2: 


Corolla blue, sky-blue, violet or lilac, brownish yellow or lilac 
whier Gry: $022........0:..0:2ae dees cote Aah cre bleh oe, 26. 
Anthers densely hairy at base and along sutures of thecae (Kopet- 
Dagyy S008. 2a et tee caee: 8. O. androssovii Novopokr. 
Anthers sparsely hairy only at base (Kara Tau) ...... AMADA SS. LSM. 
sfeitg US, seis STG Eh SR Ee Ee eles 9. O. karatavica Pavl. 
Calyx teeth deltoid at base, subulate, usually exceeding tube; corolla 
limb sky-bluish pink, but quickly losing color on drying; stem with 
few scales, often flexuous (Transcaucasia, Kopet-Dag, Pamiro-Alai) 
HRS teeing Oe ee 10. O. coelestis Boiss. and Reut. 
Calyx teeth narrowly lanceolate, usually equaling tube; corolla dif- 
ferently ‘colored)...c:ci0- nee hee et eee ia egies 2 27. 
Corolla 18-20 mm long; calyx light rae with slender teeth and 
obscure viens; inflorescence dense; stem with few ovate-lanceolate 
scales, often flexuous (Caucasus) ........ 18. O. mutelli F. Schultz. 
Corolla larger; calyx brownish, with thick teeth and prominent 
veins; stems usually with comparatively numerous, lanceolate 
SCALES, ides! sbigdelgeineate. vizssheads. dae te Pid emcee et eee Oe 28. 
Corolla (18)20—23(25) mm long; anthers glabrous or sparsely hairy 
1 eR Tid EAA eee Seon in SR GN ORNs cis I i MY. od 29. 
Corolla (23)25—30(35) mm long; anthers densely hairy at base and 
usually also along sutures Of thecae ...............:cssssesceeeeeeereeeeees Bi. 
Inflorescence lax, comparatively few-flowered, rarely dense; lon- 
gitudinal veins of corolla darker than rest of corolla ............. 
Fett GROIN ins Aone vast atv cu shall secant vast ahaa dee aes 1. O. purpurea Jacq. 
Inflorescence dense; longitudinal veins of corolla obscure ...... 30. 
Calyx teeth usually shorter than tube, subulate at tip. Parasitic on 
labiates (Transcaucasia) ..............:sesscccesreeees 13. O. cilicica G. Beck. 
Calyx teeth usually equaling tube, narrowly lanceolate. Parasitic 
on wormwood species (from Volga to Altai and Pamiro-Alai moun- 
tAlAS) ARS. ce cpce tame De ee ecguays 2. O. uralensis G. Beck. 
Inflorescence ovate, often subcapitate, dense, as long as rest of 
stem or shorter; stem usually with rather numerous scales ...... 
wisindtas sec Setaeet OM Nevhee shrarceocsrenty oe Mitng 6. O. iberica (G. Beck) Tzvel. 
Inflorescence cylindrical, rarely short-cylindrical, usually some- 


Whiat haw iokAds ORC Sy Sey Bs ee, Be a2. 
Inflorescence usually much shorter than rest of stem, stem with 
rather numerous scales. Sandy places ........... 7. O. arenaria Borkh. 


Inflorescence usually as long as rest of stem or longer; stem with 
few scales. Stony slopes (Transcaucasia) .............ccsccessecessesees 
Ee hs SO, SER, SO Seen ee 5. O. bungeana G. Beck. 


39 


33(1). 


38. 


39. 


40. 


41. 


35 


Corolla tube elongated, somewhat inflated below place of insertion 
of stamens, then narrowed and again broadened toward limb; 
stamens inserted in corolla tube away from base, often near middle 
of tube (Subsect. Inflatae G. Beck) ..............::cccsssesecssseseeeeeees 34. 
Corolla tube usually shorter in comparison with its width, broad- 
ened toward limb above place of insertion of stamens, narrowed 
below toward base, often campanulate; stamens inserted in corolla 
tube near base, rarely in its lower part (Subsect. Angustatae G. 


BEG petetisieiee srbetatin ean aphid aes eter wbaioriys deere Rv cadledn 60. 
Plant glabrous throughout ..............cccsscsseesseesesssesesesseeseeseeees 35. 
Plant somewhat glandular-hairy .................sccssssssscesssessseeseeeseees 36. 


Corolla 15-18 mm long, markedly bent forward (Baikal Region) 
eee a BB SPO patel ae cae 30. O. glaucantha Trautv. 
Corolla 18-25 mm long, mildly bent (Transcaucasia) ............... 
iuttel halons) ce Binds Beene: 50. O. glabricaulis Tzvel. 
Pubescence of inflorescence somewhat arachnoid, whitish lanate 


due} tossatherlong hairs: tz. 2nneteih.de. Darke WU eee 37. 
Pubescence of entire plant short, edlasirluter often indiscernible 
withoutimagnificationstnaht ie Secon. allow ak. 39. 
Corolla 18-23 mm long. Anthers pubescent. Parasitic on umbel- 
Dif sya denrthets oie th Peepeornd doe 2 Saas 13 ees 46. O. kotschyi Reut. 
Corolla 14-20 mm long; anthers glabrous or subglabrous. Para- 
Sitic ON WOrmWwOOd (Artemisia L.) u..........ssscccccceesessrecccceeeeeeseees 38. 


Corolla light yellow, somewhat mildly bent ........0.00 
naomi, these tne biaters bats see 4 29. O. korshinskyi Novopokr. 
Corolla blue or violet and often retaining color on drying ....... 
acuessebeel tua Soden onbe sisstssssssesessstssssseeeee 20. O, Coerulescens Steph. 
Corolla 17-25 mm long, with very narrow and long tube, sharply 
bent forward at place of insertion of stamens, then erect; anthers 
densely hairy near base and along sutures of thecae (Transbaikal 


Re GIO MS OVICE, Ran East) oncincesncaosaninancessnseandedsennctts neem 40. 
Corolla different in appearance; anthers glabrous or sparsely hairy 
dus ey. eso took unaue dern en RT LU umes anes welt Bestia. 3 41. 


Corolla light yellow, sometimes with slight bluish tinge in limb 
pevesembints. Lopeonternten lien. Formas oth fesse nel tain ys 31. O. pycnostachya Hance. 
Corolla blue or violet; inflorescence usually lax ................00 
a Ulbere un line Bas cite | 2 Ue 32. O. amurensis (G. Beck) Kom. 
Stem 8-15 cm tall, rather thick, covered with numerous, ovate, 
sometimes suborbicular scales, usually narrowed at tip into short, 
curved spine; inflorescence dense; corolla suberect. Parasitic on 
Polygonales) Wessun duianc:: 45. O. camptolepis Boiss. and Reut. 
Stem usually taller, with lanceolate or ovate-lanceolate scales. 
LOS Amt MCKIE oe. sconce apnnenananattancesbact SERS, (i008 42. 


40 


36 


42. 


49(43). 
+ 


50. 


sil 


Corolla 30-35 mm long, subglabrous, slightly bent, brownish lilac 
when dry; filaments glandular-pubescent throughout (southern 
TransCaucasia) inchs eet 37. O. sintenisii G. Beck. 
Corolla usually smaller, if large, then filaments hairy only at base 
tintnag siete shivettebadecuetamnts sank ete tal beet oen ttert wel ase sade. 43. 


. Corolla limb blue, violet or sky-blue, usually partly retaining 


color:whenrdinyes:. nese eee Rs EE SR OE, 44. 
Corolla yellow, light-yellow or brownish yellow, rarely limb with 
very faint bluish tinge, readily vanishing on drying ................ 49. 


. Corolla (20)22—35(40) mm long; stamens inserted in corolla 


tube much belowsmiddle ete eee Se. 45. 
Corolla (10)12—18(20) mm long; stamens inserted in tube usually 
slightlyabelowumiddley, .c.clux. fii82 Leek. ee 47. 


. Stamens inserted at distance of 3—5 mm from base of short and 


broad corolla tube; corolla sky-blue (Pamiro-Alai) ............... 
2 datlaan. Syaenme eet eal apanetnen on ache este ioe Sy, 36. O. clarkei Hook. 


. Corolla campanulate-tubular, limb pale violet .................... 


re ieee stein phen Gyttes cereal seueter 34. O. ariana Gontsch. 
Corolla with narrow tube and broad limb, blue or violet .......... 
(once iaal cata! nh cays ti to aha derntn cate weedy, 33. O. amoena C.A.M. 


. Corolla slightly bent forward, blue or violet, limb much broad- 


ened (Soviet Central Asia) ............:cccsseccesseeeeees 35. O. hansii Kern. 
Corolla arcuate, slightly broadened at limb ............ eee eeeeeeee 48. 


. Inflorescence usually lax, rarely dense, corolla limb bluish or 


pale violet; stem with comparatively few scales .................. 
mI eesti reas A oh as: Merb), GLb, 5 ek ATNeNy 41. O. cumana Wallr. 
Inflorescence dense; corolla limb usually blue or violet; stem 
with rather numerous, broadly lanceolate scales .................. 
BAU ach thea sabeisadstdanaaae sec Adtaueasliadaae eae eee em 40. O. cernua Loefl. 
Stamens inserted in lower 1/3 of long and suberect corolla tube; 
corolla light yellow, 20-30 mm long ..............cceeeeeeeeeeeeeeees 50. 
Stamens usually inserted slightly below middle of corolla tube; 
tube shorter compared with width and often bent forward .... 51. 
Stem up to 25 cm tall, slender, solitary, with pubescence indis- 
cemibleiwithougemasnificatiomuses. 205. 200 2 en ieee cree 
aLCTneaaigy, Bite sl euteoteney 38. O. solenanthi Novopokr. and Pissjauk. 
Stems up to 60 cm tall, often appearing in clusters, distinctly 
pubescent in upper part «2.0.0.0... cee 39. O. sulphurea Gontsch. 
Corolla 25-40 mm long, brownish yellow, distinctly bent for- 
ward))Parasitic: on umibellifers (20%. ..22008. St. HI. neon: 


41 


56. 


a7. 


+ 


60(33). 


Corolla up to 23 mm, rarely up to 25 mm long .................. 52: 
. Corolla tube arcuately bent forward; anthers glabrous or puberu- 
lentvalongtsuturcsiofsthecae sy2e.. 2a es BN RO 53: 
GarallAie be SUDEFECHON ELEC coeci sae icctctiectencvenceesainemecwecisivenwsseses 56. 


. Pubescence of inflorescence, at least on back of bract scales, 


usually comparatively long, whitish lanate. Parasitic on 
Tier |ore) | Vic C hee ene ee Ue Be Cs 46. O. kotschyi Reut. 
Pubescence of whole plant indiscernible without magnification, 
grayish or brownish. Parasitic on composites, rarely on legumes 
oo ai hie, ER LCL A MELEE LR aad BE OA os 54. 


. Corolla 20-25 mm long (Kyzyl Kum) ........ eee eeeeees 


TREE ANANSI BEY ROLE EN: 42. O. brevidens Novopokr. 
Corollasd2—20 Grimlong sek... PR SRR cee 55. 


. Plant up to 15 cm tall. Parasitic on Chesneya kopetdaghensis 


Boriss. ...........++- 44. O. badchysensis Novopokr. and V. Nikitin. 
Plant up to 35 cm tall. Parasitic on composites. ..................++ 
ie we ark hth ARE aoc A Rec 43. O. solmsii Clarke. 
Anthersidenselywhairy ati base v22.0.cnte ee ak Sik 
Anthers glabrous or sparsely hairy at base ............cceeeeeseeeees 58. 
Filaments subglabrous, inserted at distance of 5—7 mm from base 
of corolla tube (Transcaucasia). ......... 49. O. vitellina Novopokr. 
Filaments pubescent at base, inserted at distance of 7-9 mm 
from base of corolla tube (Soviet Central Asia) ................... 
Pee RIE ARIE, OI 8.5 ER eancentoes 48. O. sordida C.A.M. 


. Corolla distinctly constricted before limb. Parasitic on ivy ....... 


Pca Aebariee aaniat aed baal omer en 69. O. hederae Duby. 
Corolla tubular-campanulate, not constricted before limb ..... 59. 
Corolla with rather long, matted hairs on dorsal surface of limb. 
Parasitic mainly on umbellifers ....... 68. O. versicolor F. Schultz. 
Corolla glandular-pubescent outside, often subglabrous. Parasitic 
One R aM eS LO IE. BE 67. O. minor Sutt. 
Calyx gamosepalous, divided up to base only anteriorly (on side 
of bract scales), often 5-toothed (Caucasus) ..............cc:ccc00 61. 
Calyx divided up to base into 2 lateral, entire or bidentate seg- 
ments, sometimes connate only in front ..............eeeeeeeeeeeeee 64. 


. Calyx divided up to base anteriorly, connate on side of axis to 


considerable length. Corolla 18-22 mm long .................:e000 62. 
Calyx connate anteriorly to considerable length, usually with deep 
incisiontOn Siderofiaxis! LeU. FANN. LR Re ees 63. 


. Filaments inserted at distance of 1.5—2 mm from base of corolla 


tube, hairy only near base .............. cece 71. O. armena Tzvel. 
Filaments inserted at distance of 3-4 mm from base of corolla 
tube, hairy almost throughout .................... 71. O. armena Tzvel. 


42 


70. 


71. 


72. 


73: 


Corolla 24—35 mm long, limb dark red ... 51. O. gamosepala Reut. 
Corolla 15—18 mm long, brownish red .... 70. O. connata G. Koch. 
Corolla 23-28 mm long, sky-blue (Pamiro-Alai) ...............:eseseeeeeees 
odhsaiS sb Bulde sacle dctel Ao SRRURRL DANE. Laelia aD rae 36. O. clarkei Hook. 


Corolla.ofidifferent:colosa...c2aeeme nes. Ms. Sa 65 
Longitudinal dorsal line of corolla uniformly arcuate (Plate V, figs. 
SINT 110 0 EY ira ee enn ie Ree ce Cmaria en RO ND AN AENAEMMMPIET 39245 4.8.1. Meee 66. 
Longitudinal dorsal line of corolla erect or suberect in middle part, 
bent toward base and in limb (Plate V, figs. 6-8) ................... 78. 
Filaments hairy in basal 1/4—1/3 part, with very short, sparse, 
glandular hairs in upper part sometimes subglabrous .............. 67. 
Filaments densely hairy in basal (1/2—2/3) part, with sparse glandular 
haics iN: UPPeKPartise...csennc.snnes Nt EE, leans}. . WA. 


Stem glabrous, dark red, covered with numerous, glabrous scales. 
Parasitic on species of Salvia L. (Caucasus) ............sccesceesseeees 
siscieatesa POROONOL, see, ebb heed | ene. 81. O. colorata C. Koch. 
Stenixsomiewhat: hairy: u5, 208. 2c 68. 
Corolla campanulate, very sparsely pubescent, dark red in upper 
part of limb, yellowish in lower part. Parasitic on legumes 
(G@aucasts); 2. ecesieh.te Retest. aoe 80. O. gracilis Smith. 
Corolla tubular-campanulate, limb uniformly colored .............. 69. 
Corolla yellowish brown when dry, often with reddish or lilac 
tinge; calyx segments entire. Parasitic on species of Origanum L. 
(BatmirozAdai) ese Aloe ba soseSencswaninntn 61. O. grigorjevii Novopokr. 
Corolla yellow or brownish yellow, rarely with very slight pink 
tinge; calyx segments usually bidentate, rarely entire .............. 70. 
Corolla tube slightly bent; inflorescence usually as long as remaining 
part of stem; pubescence of stem and bract scales short, brownish. 
Parasitic ON COMPOSIEES ............:.sceeeceeceeseeeeeeeeeneeees 74. O. major L. 
Corolla tube uniformly arcuate; inflorescence usually much shorter 
than remaining part of stem; pubescence of bract scales often rather 
dense, whitish. Parasitic on umbellifers ...... 78. O. alsatica Kirschl. 
Corolla pinkish or red, often retaining color even when dry .... 72. 
Corolla yellow or brownish yellow .............::ssssscessessscseeseeeeeees aS. 
Calyx segments entire; inflorescence often ovate, corolla limb darker 
than tube. Parasitic on species of Campanula L. .............. 
acndshdiicee'sils cee EUEROsE, siaonn SR maltose” 5 ata 62. O. raddeana G. Beck. 
Calyx segments usualiy bipartite; inflorescence cylindrical; corolla 
limb lighter than pinkish tube. Parasitic on Prangos ferulacea 


hind act.cgnrls ee eth Se oom ie Bae 79. O. rosea Tzvel. 
Stem usually sparsely hairy, subglabrous. Parasitic on species of 
Thalictiumil:? sas. te ee AE 77. O. krylowii G. Beck. 


Pubescence of stem CGemserr .................ccccccssesescecccccssssccecececessssees 74. 


74. 


75. 


76. 


Th 


78(65). 
+ 


43 
Thee 


80. 


+ 


81. 
+ 


39 


Stem covered with rather numerous, long, lanceolate scales; corolla 
usually brownish yellow (Caucasus) ...........c:cccsscessscesseeeseeeees ae 
Stem with comparatively few, early dying scales; corolla usually 
light yellow A.3./4ct te, Steet oh eeee eth. ua! aesbauadea geese 11. 
Filaments hairy from base for 1/2 their length, inserted in corolla 
tube at distance of 3-4 mm from base .......... eee ceeeeeeeeeees 
winpanths elt obtener torch vonele, oe, 72. O. kurdica Boiss. and Hausskn. 
Filaments hairy from base for 2/3 length, inserted in corolla tube 


dtudistance,of2—3, mimrfrom base jibes as. dcestbondidees golbseseer. 3. dete 76. 
Inflorescence usually shorter than remaining part of stem, less 
densely covered with scales .................. 78. O. alsatica Kirschl. 


Inflorescence usually as long as rest of stem or shorter; scales of 
stem very numerous, often overlapping ..................csseeeeeeeeeee 
is, thee: cetareyaeay. dine ge eecgeem 73. O. grossheimii Novopokr. 
Stamens inserted at distance of 1.5—-3 mm from base of corolla 
tube. Parasitic on species of genus Inula L. ..........cccccccsseees 
eek) bgtectennent sls, pepercateneloses. ct 76. O. inulae Novopokr. and Abram. 
Stamens inserted at distance of 3-6 mm from base of corolla 
tube. Parasitic on species of Petasites Gaertn. and Tussillago L. 
wo. Baers titi. Behe orbiting | tetris. tells 75. O. flava Mart. 
Stamens inserted in corolla tube slightly below middle; corolla 


HO—18, coaploneyt, ..2eheod ptleviees.. bes seed. emened cgeeeenecBuk..0:s 58}. 
Stamens inserted in corolla tube much below middle, often near 
base of tube; corolla usually larger .................scesseesssesseeeeeees 79. 


Corolla 18-30 mm long, usually erecto-patent in relation to inflo- 
rescence axis, with comparatively long tube and scarcely sinuate 
galeate upper lip; longitudinal dorsal line of corolla straight for 
considerable length; calyx segments very broad, always bidentate 


peng abet alr teentbieii bined, cites bettienin, suai. sit we irenercubieis ott te ane 80. 
Corolla different in shape, much diverging from inflorescence 
axis; calyx segments entire or bidentate ...................sccceeeeeeees 84. 


Stamens inserted at distance of 1.5-3 mm, very rarely 4 mm, 
from base of corolla tube; corolla 20-30 mm long, brownish 
lilac, rarely yellowish brown, often with reddish tinge. Parasitic 
mainly on species of Galium L. ............:00000 56. O. vulgaris Poir. 
Stamens inserted at distance of 4-7 mm from base of corolla 
tube; corolla with narrower tube, 18-30 mm long ................ 81. 
Corolla brownish lilac; sos. eshte: el westitiioh esha. ane’ 82. 
Corolla yellow, limb slightly pinkish .................. es esseessesseeeeees 83. 


‘Referral back to couplet 58 is made because the species treated here, having 
stamens inserted in the middle of the comparatively narrow corolla and hence moving 
to couplet 34, belong to section Angustatae G. Beck, whose species move to couplet 


60. 


44 


40 


83. 


86. 


87. 


88. 


89. 


Calyx segments obscurely veined, usually free (Carpathian 
MOUNtAINS) <5 4.82802 LAPEER 54. O. teucrii Holand. 
Calyx segments distinctly veined, connate for considerable length 
anteriorly (from side of bract scales) (Armenia) ................... 
85). De cL EE AE 55. O. schelkovnikovii Tzvel. 
Corolla slightly inflated near base and bent forward at place of 
stamen insertion, consequently almost horizontally diverging from 
inflorescence axis (Pamiro-Alai) ...... 53. O. linczevskyi Novopokr. 
Corolla not bent at place of stamen insertion, obliquely erecto- 
patent-220 EO OBE, 52. O. lutea Baumg. 
Calyx segments broad, bidentate, connate anteriorly for consider- 
able length; corolla about 20 mm long, campanulate (Transcaucasia) 
Laclede Re, RIOR, SE 57. O. hymenocalyx Reut. 
Calyx*segmients dree, often entire - i222. ee ee 85. 
Pubescence of plant short throughout, brownish or yellowish ... 86. 
Pubescence, at least on inflorescence axis and bract scales, of longer 
whitish hairs, or plant whitish pubescent throughout (visible with 
naked eye); calyx segments entire or bidentate, subulate at tip; 
corolla without colored, dark glandular hairs ...............::ccsesseee 89. 
Calyx segments bidentate, rarely entire, pubescence of corolla 
without colored, red or violet, hairs; stamens inserted at distance of 
3-5 mm from base of corolla tube. Parasitic on composites 
(Centaurea L., Echinops L. etc.) ........::cssceeseeeeeees 74. O. major L. 
Calyx segments entire, very rarely bidentate; pubescence of corolla 
usually with red and violet glandular hairs (although in small 
number), recognized in dry material by black and dark tubercles 
(dots) at their base; filaments inserted at distance of 1-3, rarely 4 


mm, from base of corolla tube ..................00sssesssessesesseecceececeeeeees 87. 
Calyx segments in dry condition dark-colored, usually blackish, 
thicker, with ODSCUTFE VEINS .................:.ssssssssesccescccceceeecacsenessseess 88. 


Calyx segments in dry condition brownish, rarely with reddish 
tinge, distinctly veined, often markedly elongated; corolla white, 
often red or violet in limb. Parasitic on labiates .................000 
1 TE POR BON as 58. O. alba Steph. 
Corolla dark red in limb, densely covered outside with colored 
glands; stem up to 25 cm tall. Parasitic on teasel family 
(Dipsacaceae) is. .L.08.. ee. RR 60. O. reticulata Wallr. 
Corolla whitish, with pale violet or reddish tinge in limb, with few 
colored, glandular hairs outside; stem up to 60 cm tall. Parasitic ‘on 
species of Cirsium Adams .... 59. O. pallidiflora Wimm. and Grab. 
Corolla with violet or reddish tinge in limb, usually partly retained 
in dry material; inflorescence often few-flowered. Parasitic on 
Cera Trio ea Me Rd a ty 64. O. owerinii G. Beck. 


45 


41 


+ Corolla white or light yellow, rarely with pinkish or violet tinge in 
limb, quickly disappearing on drying; inflorescence usually cylin- 
deical many-flowenedyis tens. AR A eee AR 90. 

90. Corolla 20-30 mm long. Parasitic on legumes ....................:.000 
Coa, wr eilriavaniin.al meine 65. O. crenata Forsk. 

+ Corolla 16-20 mm long. Parasitic on composites ..................2+ 

ES NR cers crs a heat CAN, oak oh 66. O. picridis F. Schultz. 


Section 1. TRIONYCHON Wallroth, Schedulae crit. 1 (1822) 314; G. 
Beck, Monogr. Orob. 86 and in Pflanzenr. VI, 261 (1930) 61. Phelipaea 
sect. Trionychon C.A.M. in Ldb. FI. alt. II (1830) 459; Reuter in DC. 
Prodr. XI, 4; Ldb. Fl. Ross. III, 310; Boiss. Fl. or. IV, 495.—Calyx 
gamosepalous, campanulate, 4—5-toothed. Bracteoles present. Corolla 
with long and narrow tube, slightly broadened at place of stamen 
insertion, narrowed above, and gradually broadening toward limb; 
lower flowers in inflorescence often short-pedicellate. Stem simple or 
branched. 


Subsection 1.H o0lo0¢1ada Novopokr. in Bot. mat. gerb. Bot. 
inst. Akad. Nauk SSSR, XIII, 1(50)305, pro sect.—Stem simple, usually 
with comparatively numerous lanceolate scales. Perennials, rarely 
biennials. 


1. O. purpurea Jacq. Enum. agri Vindob. (1762) 108 and Obs. 
(1762) 252; G. Beck, Monogr. Orob. (1890) 123, tab. II, f. 28 and in 
Pflanzenr. IV, 261 (1930) 105; Fedch. and Fler. Fl. 890, fig. 857; 
Grossh. Fl. Kavk. IV, 10.—0O. coerulea Vill. Hist. pl. Dauph. II (1787) 
406; M.B. FI. taur.-cauc. II, 82 and III, 417; Schmalh. Fl. I, 294.— 
? O. pareysii G. Beck in Oesterr. bot. Zeitschr. (1884) 310.— Phelipaea 
coerulea C.A.M. Verzeichn. Pfl. Cauc. (1831) 104; Reuter in DC. 
Prodr. XI, 5; Ldb. Fl. Ross. III, 311; Boiss. Fl. or. IV, 495.—Ic.: 
Rchb. Ic. Fl. Germ. XX, tab. 149; G. Beck, l.c.; Fedch. and Fler. FI. 
Evrop. Ross.; Hegi, III. Fl. VI, f. 81 (a-d). 

Perennial. Plant 15-50 cm tall, glandular-puberulent (“mealy”). 
Stem yellow or brownish, usually with reddish tinge, covered with 
lanceolate, acute, 10-15 mm long scales. Inflorescence usually short- 
cylindrical, up to 10-15 cm long, lax and often comparatively few- 
flowered, rarely dense, usually much shorter than remaining part of 
stem. Bract scales 8-12 mm long, lanceolate or ovate-lanceolate, like 
bracteoles usually reaching only up to base of calyx teeth, glandular- 
puberulent on back and along margin. Bracteoles linear-subulate. 
Calyx 8-14 mm long, with very prominent and darker veins, glandular- 
puberulent; lateral calyx teeth deltoid at base, narrowly lanceolate, 


46 


42 


somewhat equaling tube. Corolla (18)20—23(25) mm long, lilac or blu- 
ish, paler toward base, with darker longitudinal veins, glandular-pu- 
berulent outside, glabrous inside, excepting densely hairy folds of lower 
lip; broadly ovate, sometimes subacute, subcrenate. Filaments glabrous 
or hairy at base; anthers glabrous or sparsely hairy. May to July. 

Parasitic on composites, mainly on species of Achillea L.; also 
reported on species of Artemisia L., Pyrethrum Gaertn., Anthemis L. 
Steppes, stony slopes, scrub.—European USSR: Upper Dniester, 
Middle Dnieper, Volga-Don, Trans-Volga Region, (south-west), 
Bessarabia, Black Sea Region, Lower Don, Lower Volga, Crimea. 
Caucasus: All regions. General distribution: Atlantic and Central 
Europe, Mediterranean Region, Balkans-Asia Minor, Armenia- 
Kurdistan, Iran. Described from Austria. Type in Vienna. 

Note. Among the varieties of this species cited by G. Beck, we 
consider var. iberica G. Beck as a separate species, closer to O. 
arenaria Borkh., and assign the specimen from Dagestan identified by 
Beck as var. simulans G. Beck to the latter species. The third variety, 
var. pareysii G. Beck [Monogr. Orob. (1890) 310 (= O. pareysii G. 
Beck)], characterized by a taller stem, branched in the middle, dense 
and many-flowered inflorescence and corolla up to 30 mm long, 
apparently is described from random, not fully typical specimens, 
possibly also belonging to O. arenaria Borkh. Beck reported it for 
Crimea and Talysh. 


2. O. uralensis G. Beck, Monogr. Orob. (1890) 132 and in 
Pflanzenr. IV, 261 (1930) 112.—Phelipaea pallens Bge. in Ldb. FI. 
Ross. III, 312; Korsh. Tentam, fl. Ross. or. 327; O. and B. Fedch. 
Perech. rast. Turkest. V, 116, non Orbanche pallens F. Schultz, 1840. 

Perennial. Plant 15—40 cm tall, glandular-pubescent. Stem yellow- 
ish brown, sometimes with reddish tinge; 3-5 mm thick in middle, 
covered with lanceolate, acute, up to 15 mm long scales. Inflorescence 
ovate or short-cylindrical, dense, usually much shorter than remaining 
part of stem. Bract scales 8-12 mm long, ovate-lanceolate, almost 
equaling calyx, glandular-pubescent. Bracteoles linear-subulate, slightly 
shorter than calyx. Calyx 9-12 mm long, with very prominent veins, 
glandular-pubescent; lateral teeth deltoid at base, narrowly lanceolate, 
almost equaling tube. Corolla 18-25 mm long, lilac, bluish or pale 
violet, lighter, whitish at base of tube, with obscure longitudinal veins, 
glandular-pubescent outside, subglabrous inside, excepting densely hairy 
folds of lower lip; lobes of lower lip broadly ovate, sometimes sub- 
acute, ciliolate along margin, subcrenate. Filaments sparsely pubescent 
at base, glabrous above; anthers somewhat hairy at base. Style glandu- 
lar-puberulent near stigma. June to August. 


47 


43 


Parasitic on species of wormwood (Artemisia latifolia Ldb., A. 
armeniaca Lam., A. dracunculus L., A. glauca Pall., A. frigida Willd., 
A. sieversiana Willd., etc., less commonly on other species). Steppes, 
stony slopes, scrub.— European USSR: Volga-Kama (south-western 
part), Volga-Don (eastern part), Trans-Volga region, Lower Volga 
(north-eastern part). Western Siberia: Upper Tobol, Irtysh (eastern 
part), Altai Mountains. Soviet Central Asia: Aralo-Caspian Region 
(northern part), Balkhash Region (eastern part), Dzhungaria-Tarbagatai, 
Tien Shan, Pamiro-Alai, Syr Darya, mountainous Turkmenia. General 
distribution: Iran (?), Dzhungaria-Kashgaria. Described from the area 
of Lake Inder. Type in Paris (?). 

Note. This species occupies an intermediate position between O. 
purpurea Jacq. and O. caesia Rchb., and, apparently, splits into several 
physiological and partly also geographical races, differing slightly 
morphologically and approaching one or the other of these species. 
Described by Bunge from a random and atypical sample, with 6-toothed 
calyx. 


3. O. caesia Rchb. pat. Iconogr. VII (1829) 48, f. 936; Reut. in 
DC. Prodr. XI, 6, p.p.; G. Beck, Monogr. Orob. 120, tab. II, f. 27 and 
in Pflanzenr. IV, 261, 112; Fedch. and Fler. Fl. 840, fig. 836; Grossh. 
Fl. Kavk. IV, 10; Kryl. Fl. Zap. Sib. X, 2537.—O. asiatica Weinm. 
in Bull. Soc. Nat. Mosc. XXIII (1850) 552.—O. lanuginosa G. Beck 
apud Kryl in Tr. Obsch. estestv. Kazansk. univ. (1881) 202; Schmalh. 
Fl. II, 293.—Phelipaea lanuginosa C.A.M. in Ldb. Fl. Alt. II (1830) 
460; Reut. I.c. 4; Ldb. Fl. Ross. III, 310; Boiss. Fl. or. [V, 496.—Ie.: 
Rchb. I.c.; Ldb. Ic. pl. fl. ross. IV, tab. 337; G. Beck, l.c. 

Perennial or biennial. Plant 10-35 cm tall, arachnoid-villous with 
rather long, white hairs in inflorescence elsewhere glandular-pubes- 
cent. Stem yellowish brown, often with reddish tinge, 3-5 mm thick 
in middle, slightly thickened at base, covered with lanceolate or ovate- 
lanceolate, 8-15 mm long scales. Inflorescence ovate or short-cylin- 
drical, dense, rarely somewhat lax, usually much shorter than remain- 
ing part of stem. Bract scales 8-14 mm long, ovate-lanceolate, almost 
equaling calyx, somewhat arachnoid-villous with white hairs at back 
and along margin. Bracteoles linear-subulate, slightly shorter than 
calyx, usually arachnoid-lanate with white hairs. Calyx 9-14 mm long, 
with prominent veins, glandular-hairy, often somewhat whitish 
pubescent; lateral teeth narrowly lanceolate, almost equaling tube. 
Corolla 18-25 mm long, usually distinctly bent forward, sky-blue, 
bluish or pale violet, lighter at base of tube, with obscure longitudinal 
veins, glandular-hairy outside, with mixture of comparatively long 
hairs, glabrous inside, excepting densely hairy folds and sparsely 


48 


44 


pubescent lobes of limb; lobes of lower lip broadly ovate, obtuse, 
sometimes subacute, crenate and ciliate along margin. Filaments 
sparsely pubescent or glabrous at base, glabrous above; anthers 
glabrous, rarely sparsely hairy at base. Style glandular-puberulent near 
stigma. June to August. 

Parasitic on species of wormwood (Artemisia austriaca Jacq., A. 
maritima L. s. 1. etc.). Steppes, stony slopes, old fields.— European 
USSR: Volga-Kama, Upper Dnieper (southern part), Volga-Don, Trans- 
Volga Region, Middle Dnieper, Bessarabia, Black Sea Region, Lower 
Don, Lower Volga, Crimea. Caucasus: Caucasian foothills, Dagestan, 
eastern and southern Transcaucasia, Talysh. Siberia: Ob’ inlet (southern 
part), Upper Tobol, Irtysh, Altai Mountains. Eastern Siberia: Angara- 
Sayan. Soviet Central Asia: Aralo-Caspian Region, Balkhash Region, 
Dzhungaria-Tarbagatai, Tien Shan, Syr Darya, Pamiro-Alai (?). General 
distribution: Central Europe (southeastern part), Balkans-Asia Minor, 
Armenia-Kurdistan, Iran, Dzhungaria-Kashgaria. Described from Aus- 
tria. Type in Vienna. 

Note. In the USSR, the morphologically poorly defined var. bo- 
realis G. Beck. [Monogr. Orob. (1890) 122 (= Phelipaea lanuginosa 
C.A.M.)], with a larger (21-25 mm long) and less curved corolla 
predominates. The name “borealis” was taken by G. Beck from the 
species O. borealis Turcz. (= O. coerulescens Steph.), described by 
Turczaninov, which Beck mistakenly considered close to O. caesia 
Rchb. 


4. O. pulchella (C.A.M.) Novopokr. in Grossh. Opred. rast. Kavk. 
(1949) 354; Sosnovsk. in Fl. Gruzii, VII, 644.—O. aegyptiaca f. 
pulchella G. Beck, Monogr. Orob. (1890) 102 and in Pflanzenr. (1930) 
84.—Phelipaea pulchella C.A.M. in Eichw. Pl. casp.-cauc. (1831) 
17, tab. XVIII; Reuter in DC. Prodr. XI, 5; Ldb. Fl. Ross. III, 311; 
Boiss. Fl. or. IV, 496.—Ic.: C.A.M. l.c. 

Perennial. Plant 8—15 cm, rarely 25 cm, tall, glandular-hairy, some- 
what arachnoid-villous with white hairs in inflorescence. Stem yel- 
lowish brown, usually with red or violet tinge, slender and slightly 
thickened at base, 2—4 mm thick in middle, covered with lanceolate or 
ovate-lanceolate, 8-10 mm long scales. Inflorescence ovate, often 
subcapitate, rarely short-cylindrical, comparatively few-flowered, dense 
or somewhat lax, usually shorter than remaining part of stem, rarely 
almost equaling it. Bract scales ovate-lanceolate, up to 10 mm long, 
almost equaling calyx, somewhat arachnoid-villous with white hairs at 
back and along margin. Bracteoles linear-subulate, slightly shorter 
than calyx, usually arachnoid-lanate. Calyx 9-12 mm long, obscurely 
veined, somewhat lanate; lateral teeth narrowly lanceolate, sometimes 


49 


45 


subulate at tip and almost equaling tube. Corolla (20)22—27(30) mm 
long, suberect, blue, lighter at base of tube, with dense and rather long 
hairs outside (often shortly grayish-lanate throughout), glabrous inside, 
excepting densely hairy folds and sparsely hairy lobes of limb; lobes 
ciliate and crenate along margin; lobes of lower lip orbicular or broadly 
ovate, sometimes subacute. Filaments pubescent at base, glabrous above 
or sparsely glandular-hairy near anthers; anthers densely hairy at base, 
and often also along sutures of thecae. Style sparsely glandular-hairy 
near stigma. June to July (Plate IV, fig. 1). 

Parasitic on Elisanthe noctiflora (L.) Rupr.; also reported on 
Teucrium chamaedrys L. and Galium verum L. Stony slopes, mainly 
in middle and lower mountain zones.— Caucasus: Western, southern 
and eastern Transcaucasia. General distribution: Asia Minor, Armenia- 
Kurdistan. Described from southern Georgia. Type in Leningrad. 

Note. In the Megri District of the Armenian SSR, this species was 
collected twice on the roots of Elisanthe noctiflora (L.) Rupr. How- 
ever, it is possible that plants from the vicinity of Tbilisi and from the 
extreme south of Georgia have other host plants and represent distinct 
physiological races. 


5. O. bungeana G. Beck, Monogr. Orob. (1890) 119, tab. II, f. 25 
and in Pflanzenr. IV, 261 (1930) 97; Grossh. Fl. Kavk. IV, 9.—O. 
caucasica G. Beck in Fedde, Repert, XVIII (1922) 34 and in Pflanzenr., 
IV, 261 (1930) 101; Grossh. Fl. Kavk. IV, 10.—Ic.: G. Beck, l.c. 
(1890). 

Perennial. Plant 15-30 cm tall, usually densely glandular-pubes- 
cent, rarely with longer hairs, whitish-lanate. Stem yellowish brown, 
sometimes with reddish tinge, often flexuous, 3-5 mm thick in middle, 
usually with few caducous scales. Inflorescence usually short-cylin- 
drical, somewhat lax rarely dense, usually longer than rest of stem or 
equaling it, up to 18 cm long. Bract scales ovate-lanceolate, 8-14 mm 
long, usually slightly shorter than calyx, densely glandular-hairy, some- 
times shortly whitish-lanate. Bracteoles linear-subulate, slightly shorter 
than calyx. Calyx 12-15 mm long, with very prominent veins, usually 
grayish due to dense glandular pubescence; lateral teeth narrowly lan- 
ceolate, often subulate at tip, equaling calyx tube or slightly longer. 
Corolla (22)25-30(35) mm long, markedly, but gradually broadened 
toward limb, suberect, bluish violet, lighter toward base of tube, densely 
glandular-pubescent outside, glabrous inside, excepting densely hairy 
folds and sparsely hairy lobes of limb; limb ciliate and subcrenate 
along margin; lobes of lower lip broadly ovate, sometimes subacute. 
Filaments diffusely hairy at base, glabrous above; anthers densely 


50 


46 


hairy at base, and often along sutures of thecae. Style sparsely glandular- 
pubescent. May to June. 

Parasitic on Gundelia tournefortii L., possibly aiso on other com- 
posites. Stony and rubbly slopes of lower mountain zone. — Caucasus: 
Southern and eastern Transcaucasia. General distribution: Armenia- 
Kurdistan, Iran-Afghanistan (western part). Described from northern 
Iran. Type in Leningrad. . 

Note. Although the type specimen of O. bungeana G. Beck shows 
general similarity to O. pulchella (C.A.M.) Novopokr. (small size of 
plant, subcapitate inflorescence, similar color of flower), the calyx 
and corolla of this species show greater similarity to O. caucasica G. 
Beck, described later also by G. Beck. Therefore, we considered it 
possible to combine both these species, since G. Beck, with only very 
inadequate material from the Caucasus, described large and small plants 
of the same species as different species. We also could not agree with 
I.V. Novopokrovsky who, in “Opredelitel Rast. Kavkaza” of A.A. 
Grossheim (1949), treats O. bungeana G. Beck as a separate species 
and considers O. caucasica G. Beck as synonym of O. pulchella 
(C.A.M.) Novopokr. However, it should be noted that O. bungeana G. 
Beck, as it is treated here, is not an entirely homogeneous species, 
varying firstly in the degree of pubescence of the inflorescence and, 
perhaps, comprising two or three physiological races. 


6. QO. iberica (G. Beck) Tzvel. comb. n.—O. purpurea var. iberica 
G. Beck, Monogr. Orob. (1890) 126 and in Pflanzenr. IV, 261 (1930) 
111; Grossh. Fl. Kavk. IV, 10. 

Perennial. Plant 10-20 cm tall, densely glandular-pubescent. Stem 
yellowish brown, 3—5 mm thick in middle, slightly thickened at base, 
covered with rather numerous, ovate-lanceolate, up to 14 mm long 
scales. Inflorescence ovate, often subcapitate, dense, as long as re- 
maining part of stem or shorter. Bract scales ovate-lanceolate, up to 
14 mm long, slightly shorter than calyx, glandular-pubescent at back 
and along margin. Bracteoles linear-subulate, slightly shorter than 
calyx. Calyx 8-14 mm long, light brown, with slightly prominent 
viens, but rather thick; lateral teeth narrowly lanceolate, almost equaling 
calyx tube, glandular-pubescent. Corolla (23)25—30(32) mm long, 
markedly, but gradually broadened toward limb, suberect, lilac or blue 
(?), often dull pink when dry, sparsely glandular-puescent outside, 
glabrous inside, excepting densely hairy folds and sparsely hairy lobes 
of limb; limb ciliate and subcrenate along margin; lobes of lower lip 
orbicular or broadly ovate, sometimes subacute. Filaments sparsely 
hairy at base, glabrous above or sparsely glandular-puberulent under 


53 


47 


anthers; anthers densely hairy at base, and usually also along sutures 
of thecae. Styie glandular-puberulent near stigma. May to June. 

Parasitic on species of wormwood (Artemisia meyeriana Bess. 
and others). Steppe, dry slopes.— Caucasus: Eastern Transcaucasia, 
southern Transcaucasia (?). General distribution: Asia Minor, Armenia- 
Kurdistan. Described from Turkey. Type in Vienna. 

Note. This species was treated by G. Beck as a variety of O. 
purpurea, although in many essential features (longer pubescence, 
large corolla, densely hairy anthers) it is, in our opinion, much closer 
to O. arenaria Borkh. and undoubtedly represents a separate race, 
widely distributed in wormwood semideserts of eastern Azerbaidzhan. 


7. O. arenaria Borkh. in Romer’s Neuem Mag. f. Bot. I (1797) 
6; G. Beck, Monogr. Orob. 129, tab. I, f. 29 and in Pflanzenr. IV, 
261 (1930) 105, f. 12 (B—L); Schmalh. FI. II, 294; Fedch. and Fler. 
Fl. 891, fig. 858; Grossh. Fl. Kavk. IV, 10.—O. laevis L. Sp. pl. 
(1753) 632, p.p. [momen ambiguum, vide G. Beck in Vierteljahrschr. 
nat. Ges. Ziirich, 71 (1902) 162]; Goncharov vo. Fl. Yugo-Vost. VI, 
236.—O. robusta Dietr. Fl. boruss. III (1835) tab. 151.—Phelipaea 
arenaria Walp. Repert. bot. III (1844) 459; Reut. in DC. Prodr. XI, 
6; Ldb. Fl. Ross. III, 312; Boiss. Fl. or. IV, 495.—P. robusta Walp. 
l.c. 460; Reut l.c. 9; Ldb. I.c. 313.—Ie.: Rchb. Ic. Fl. Germ. XX, tab. 
145 and 146; Dietr. l.c.; G. Beck, l.c.; Fedch. and Fler. 1.c.; Hegi, Ill. 
Fl. VI, f. 81 (e-i). 

Biennial or. perennial. Plant 15-50 cm tall, abundantly glandular- 
pubescent. Stem yellowish brown, often with reddish tint, 3-6 mm 
thick in the middle part, slightly thickened at base, occasionally 
branched in inflorescence, bearing fairly numerous lanceolate, up to 
20 mm long, scales. Inflorescence cylindrical, rarely short cylindrical 
or oval, lax or fairly thick, multiflowered, usually considerably shorter 
than remaining part of stem. Bract scales lanceolate, gradually taper- 
ing, up to 20 mm long, nearly equaling calyx, densely glandular- 
pubescent on dorsum and along margin. Bracteoles linear-subulate, 
thick, usually reaching only to base of calyx teeth. Calyx 10-18 mm 
long, light brown, sometimes densely glandular-pubescent and hence 
grayish, with prominent veins, lateral teeth deltoid at base to narrowly 
lanceolate, often subulate at tip, usually somewhat longer than calyx 
tube. Corolla 25-35 mm long, considerably broadened at limb, suberect, 
lilac or blue, paler at base of tube, glandular-pubescent outside, gla- 
brous inside, besides abundantly hairy folds and sparsely hairy lobes 
of limb; margin of lobes fimbriate and weakly crenate; lobes of lower 
lip orbicular or broadly ovate, occasionally acute. Filaments sparsely 
puberulent at base; anthers densely hairy at base and usually at suture 
of thecae. Style glandular-puberulent below stigma. May to June. 


48 


51 


Plate IV. 
1. Orobanche pulchella (C.A.M.) Novopokr., general appearance of plant, 
flower. —2. O. coelestis Boiss. and Reut., flower.—3. O. kelleri Novopokr., 
flower. —4. O. septemloba (G. Beck) Tzvel., flower.—5. O. aegyptiaca Perts., 
flower.—6. O. sogdiana Novopokr., general appearance of plant, flower. — 
7. O. ramosa L., flower. 


54 


49 


Parasitic on species of Artemisia (A. marschalliana Spreng., A. 
campestris L. and others); also reported on Centaurea arenaria M.B. 
Steppe, slopes, scrub; predominantly in sandy soils.— European USSR: 
Upper Dnieper, Middle Dnieper, Volga-Don, Bessarabia, Black Sea, 
Lower Don, Lower Volga, Crimea. Caucasus: Ciscaucasia, Dagestan, 
eastern Transcaucasia, southern Transcaucasia(?), Talysh. Western 
Siberia: Upper Tobol (southwestern part). Soviet Central Asia: Aralo- 
Caspian (northwestern part). General distribution: Central and Atlantic 
coast of Europe, Mediterranean, Balkans-Asia Minor. Described from 
Germany. Type in Berlin (?). 

Note. Plants with branched stems occasionally occur —f. robusta 
(Dietr.) G. Beck [Monogr. Orob. (1890) 130 (= O. robusta Dietr.)]. 
We also refer to O. arenaria Borkh. the plants identified by Beck as 
O. purpurea var. simulans G. Beck (l.c. 127). 


8. O. androssovii Novopokr. in Bot. mat. Gerb. Bot. inst. AN 
SSSR, XIII (1950) 309. 

Perennial. Plant up to 20 cm tall, glandular-pubescent, almost 
glabrous below. Stem yellowish brown, 3-5 mm thick in middle part, 
slightly thicker at base, often sinuate and emerging in bunches of 2 or 
3 from the root of the host plant, densely covered with 1—1.2 cm long 
lanceolate scales. Inflorescence cylindrical or shortly cylindrical, thick, 
more lax below, multiflowered, more or less equaling remaining part 
of stem, 8-11 cm long. Bract scales ovate-lanceolate, 8-12 mm long, 
usually somewhat shorter than calyx, glandular-pubescent. Bracteoles 
linear-subulate, somewhat shorter than calyx. Calyx 9-11 mm long, 
light brown, with slightly prominent darker veins, glandular-pubes- 
cent, lateral teeth narrowly lanceolate, shortly subulate at tip, equaling 
or slightly longer than calyx tube. Corolla 20-25 mm long, suberect, 
light blue, light yellow when dry, glandular-pubescent outside, gla- 
brous inside, moreover folds and lobes of limb densely hairy, margin 
of the lobes fimbriate and weakly crenate; lobes of lower lip orbicular 
or broadly ovate, obtuse. Filaments sparsely pubescent at base, gla- 
brous above or glandular-pubescent near anthers; anthers densely hairy 
at base and along sutures of thecae. Style sparsely glandular-pubes- 
cent. May to June. 

Host plant unknown; apparently on species of Artemisia L. Rubbly 
slopes of lower mountain belt.— Soviet Central Asia: mountainous 
Turkmenia. General distribution: probably Iran. Described from the 
environs of Firyuza. Type in Leningrad. 


9.0. Karatavica Pavl. in Vestn. AN Kazakhsk. SSR, 4 (1951) 94. 


50 


Perennial. plant 15—25 cm tall, glandular-pubescent, almost gla- 
brous below. Stem yellowish, in the middle part 3-6 mm thick, nearly 
as thick at base, covered with 9-13 mm long lanceolate scales. Inflo- 
rescence shortly cylindrical, dense, multiflowered, more or less equal 
to remaining part of stem, 5-10 cm long. Bract scales lanceolate, 10— 
13 mm long, equaling the calyx or slightly shorter, glandular-pubes- 
cent. Bracteoles linear-subulate, usually reaching only up to base of 
calyx teeth. Calyx 11-13 mm long, yellowish, with mildly prominent 
veins, sparsely glandular-pubescent, teeth lanceolate at base, nearly 
equaling calyx tube. Corolla 18-20 mm long, suberect, whitish or 
light blue in limb, light yellow when dry, sparsely glandular-pubescent 
outside, glabrous inside, folds of lower lip densely hairy, lobes of 
limb sparsely pubescent, lobes of lower lip orbicular or broadly ovate, 
obtuse, fimbriate or weakly crenate along margin. Filaments glabrous 
or sparsely hairy at base; anthers sparsely hairy at base. Style glandular- 
pubescent under stigma. June to July. 

Parasitic on Artemisia karatavica Krasch. and Abol. Rubbly 
slopes. — Soviet Central Asia: Tien Shan (Kara Tau Range). Endemic. 
Described from Kazakhstan. Type in Alma-Ata, cotype in Leningrad. 


10. O. coelestis Boiss. and Reut. in Pinard, Pl. exs. (1843); G. 
Beck, Monogr. Orob. 114, tab. II, f. 21 and in Pflanzenr. IV, 261, 93; 
Grossh. Fl. Kavk. IV, 10 and Opred. rast. Kavk. (1949) 354.—O. 
persica G. Beck in herb.; Novopokr. in Grossh. Opred. rast. Kavk. 
(1949) 354.—Phelipaea coelestis (Boiss. and Reut.) Reuter in DC. 
Prodr. XI, 5; Boiss. Fl. or. 1V, 496—P. tricholoba var. simplex Reuter, 
lic. 10.—P. syspirensis C. Koch in Linnaea, XXII (1849) 662.—Ic.: 
G. Beck, Monogr. Orob. 

Biennial or perennial (?). Plant up to 30 cm tall, glandular-pubes- 
cent, nearly glabrous below. Stem yellowish, in middle part 2-7 mm 
thick, slightly thickened at base, often sinuate, sparsely covered (rarely 
densely) with ovate-lanceolate or lanceolate, 8-15 mm long, scales. 
Inflorescence cylindrical or oval (in small-sized specimens nearly capi- 
tate), dense and multiflowered, longer than remaining part of stem, 
rarely almost equaling stem, up to 20 cm long. Bract scales lanceolate, 
gradually acuminate, 8-18 mm long, usually not reaching tip of calyx 
teeth, glandular-pubescent. Bracteoles lanceolate-subulate with thick- 
ened middle veins, often arcuate when dry, shorter than calyx. Calyx 
10-15 mm long, light brown, with mildly prominent brownish veins, 
glandular-pubescent, lateral teeth attenuated, subulate with lanceolate 
base, usually exceeding calyx tube three and a half times, rarely almost 
equaling, usually deflexed sideways in fruit, incised 2-2.5 mm from 
the side of the inflorescence axis, not reaching base of calyx, 5th 


56 


51 


tooth usually short. Corolla 18-20 mm long, mildly bent forward, 
whitish with bluish-pinkish limb, often entirely yellowish or pinkish 
whey dry, densely glandular-puberulent outside, densely hairy inside 
and on folds of lower lip, lobes of limb fimbriate and mildly crenate 
along margin, upper lip wide, semicircular, lower broadly ovate, obtuse, 
rarely subacute. Filaments nearly glabrous, with few short hairs at 
base; anthers sparsely hairy. Style glandular-puberulent under stigma. 
May to June (Plate IV, fig. 2). 

Parasitic on Cousinia Cass. and Centaurea L. Also reported on 
Eryngium L. Rubbly and clayey slopes of lower and middle mountain 
belts. — Caucasus: Southern Transcaucasia, Talysh. Soviet Central Asia: 
mountainous Turkmenia, Pamiro-Alai. General distribution: Eastern 
Mediterranean, Asia Minor, Armenia-Kurdistan, Iran, Dzhungaria- 
Kashgaria (southern part). Described from Turkey. Type in Geneva. 

Note. The larger plants of this species have long inflorescences 
with extremely long calyx teeth and were referred by Beck to O. 
coelestis Boiss. et Reut. f. persica G. Beck. (l.c. (1890) 115). The 
Herbarium of the Botanical Institute of the Academy of Sciences of 
the USSR possesses extensive material which convincingly shows that 
this form has no taxonomic significance. O. heldreichii (Reut.) G. 
Beck was reported from the Caucasus on the basis of one poor specimen 
of O. coelestis from Lake Sevan, which was misidentified by Beck. 
Apparently, this species occurs only in Turkey. 


11. O. borissovae Novopokr. in Bot. mat. Gerb. Bot. inst. AN 
SSSR, XIII (1950) 305. 

Biennial or Perennial. Plant up to 35 cm tall, glandular-hairy, 
white-lanate or villous in inflorescence. Stem yellowish, 3-5 mm thick 
in middle part, slightly thicker at base, usually with few ovate- 
lanceolate scales. Inflorescence cylindrical, dense, multiflowered, 
slightly longer than remaining part of stem or equaling it. Bract scales 
lanceolate, gradually acuminate, 12-15 mm long, usually not reaching 
tip of calyx teeth, more or less whitish-lanate on back. Bracteoles 
linear-subulate, slightly shorter than calyx. Calyx 14-16 mm long, 
light brown, with slightly prominent veins, densely glandular-hairy, 
teeth subulate with lanceolate base often exceeding calyx tube two 
and a half times. Corolla 18-23 mm long, bent forward slightly, light 
blue (?), densely glandular-hairy outside, inside densely hairy only on 
folds of lower lip. Lobes of limb fimbriate and mildly crenate along 
margin, upper lip semi-orbicular, lower lip broadly ovate, obtuse. 
Filaments nearly glabrous, with few hairs at base; anthers densely 
hairy at base and usually along sutures of thecae. Style densely 
glandular-puberulent. June to July. 


37) 


52 


Host plant not precisely known; reported on Artemisia sp. Rubbly 
slopes. —Soviet Central Asia: mountainous Turkmenia. General dis- 
tribution: Iran. Described from Kopet Dag. Type in Leningrad. 


12. O. kelleri Novopokr. in Bot. mat. Gerb. Bot. inst. AN SSSR, 
XIII (1950) 908. 

Biennial or perennial (?). Plant 10-25 cm tall, glandular-pubes- 
cent, base often nearly glabrous. Stem yellowish brown, 3-8 mm thick 
in middle part, slightly thicker at base with relatively few deltoid- 
ovate or broadly lanceolate, up to 8-10 mm long, scales. Inflorescence 
short cylindrical, rarely cylindrical or oval, fairly dense and 
multiflowered, nearly equal in length to remaining part of stem, up to 
14 cm long. Bract scales ovate, upper sometimes attenuated into short 
lanceolate apex, short, 6-9 mm long, nearly equaling calyx, glandular- 
pubescent. Bracteoles linear-subulate, shorter than calyx. Calyx 8-11 
mm long, somewhat inflated in fruit, light brown, with slightly notice- 
able veins, glandular-pubescent teeth deltoid at base, short, subulate, 
often recurved in fruit usually shorter than calyx tube (sometimes 
nearly twice as long). Corolla 18—20 mm long, relatively less broadened 
in limb, clearly bent forward, whitish, with bluish limb, often entirely 
yellowish when dry, glandular-pubescent outside, glabrous inside (par- 
ticularly folds of lower lip), rarely with solitary hairs on lobes of limb 
and in front portion of folds; lobes of limb with fimbriate and nearly 
entire margin, slightly recurved, lower lip broadly ovate, obtuse. Fila- 
ments glabrous or with few short hairs at base; anthers mildly hairy 
at base, nearly glabrous. Style glabrous or subglabrous. May to June 
(Plate IV, fig. 3). 

Parasitic on Kochia prostrata (L.) Schrad. Clayey and rubbly 
slopes, sands.—Caucasus: Eastern and southern Transcaucasia. Western 
Siberia: Altai Range (southern part). Soviet Central Asia: Aralo- 
Caspian, Balkhash Region. General distribution: probably Armenia- 
Kurdistan and Dzhungaria-Kashgaria. Described from eastern 
Kazakhstan. Type in Leningrad. 

Note. Despite considerable gaps in the distribution (but, probably, 
only appearing so because of insufficient herbarium material) the 
identity of specimens of this species from southern Transcaucasia and 
Kazakhstan is not to be doubted. Moreover, this is further substanti- 
ated by the fact that the species is parasitic on the same host plant. 


13. O. cilicica G. Beck, Monogr. Orob. (1890) 119, tab. II, f. 26 
and in Pflanzenr. IV, 261 (1930) 96.—Ic.: G. Beck, l.c. 

Biennial or perennial (?). Plant 8-20 cm tall, densely glandular- 
pubescent, base sometimes subglabrous. Stem yellowish brown, middle 


58 


53 


part 3-6 mm thick, slightly thicker at base, with relatively few deltoid- 
ovate or lanceolate, up to 8-10 mm long, scales. Inflorescence oval or 
short cylindrical, fairly dense and multiflowered, almost equaling re- 
maining part of stem or shorter, up to 8 cm long. Bract scales broadly 
lanceolate, often acuminate, 6-10 mm long, equaling calyx or slightly 
shorter, densely glandular-pubescent. Bracteoles linear-subulate, shorter 
than calyx. Calyx 8-14 mm long, somewhat inflated in fruit, light 
brown, with slightly noticeable veins, glandular-pubescent, its teeth 
deltoid at base to lanceolate, subulate at tip, often reflexed in fruit, 
shorter than or nearly equaling calyx tube. Corolla 18-24 mm long, 
dilated close to base towards limb, distinctly bent forward, pale blue, 
lighter in color towards base, often entirely brownish yellow when 
dry, glandular-pubescent outside, subglabrous inside, but folds of lower 
lip glandular-pubescent, lobes of limb with fimbriate, entire or weakly 
crenate margin, lower lip broadly ovate, obtuselike. Filaments with 
few glandular hairs; anthers sparsely hairy at base. Style rarely 
glandular-pubescent. June to July. 

Parasitic on Stachys inflata Benth.; also reported on species of the 
genera Salvia L. and Phlomis L. Rubbly slopes of lower mountain 
belts. — Caucasus: Southern and eastern (?)Transcaucasia. General 
distribution: Asia Minor, Armenia-Kurdistan, Iran. Described from 
Turkey. Type in Vienna. 

Note. Evidently in the Caucasus there are physiological races close 
to this species parasitizing members of the Labiatae. Thus, in the 
Herbarium of the Botanical Institute of the Academy of Sciences of 
the USSR, there are specimens from Dagestan closely resembling O. 
cilicica G. Beck which we, for the time being, prefer to assign to O. 
mutelii F. Schultz var. spissa (Rouy) G. Beck. 


14. O. septemloba (G. Beck) Tzvel. comb. n.—O. oxyloba var. 
septemloba G. Beck in Fedde. Repert. XVIII (1922) 34 (pro forma) 
and in Pflanzenr. IV, 261 (1930) 90. 

Biennial or perennial (?). Plant up to 25 cm tall, glandular-pubes- 
cent. Stem yellowish brown, middle part 3—5 mm thick, slightly thicker 
at base, bearing fairly numerous lanceolate, 7-12 mm long, scales. 
Inflorescence cylindrical, rarely oval, dense and multiflowered, as long 
as or longer than remaining part of stem or up to 18 cm long. Bract 
scales lanceolate, 6-10 mm long, usually not reaching upper teeth of 
calyx, glandular-pubescent. Bracteoles linear-subulate, slightly shorter 
than calyx. Calyx 7-8 mm long, sometimes slightly inflated in fruit, 
light brown, with faint veins, glandular-pubescent, teeth lanceolate- 
subulate, reflexed in fruit, usually somewhat longer than calyx tube. 
Corolla 16-20 mm long, somewhat inflated below the point of inser- 


59 


54 


tion of stamen, tapered above and from this point gradually and slightly 
dilated towards limb, distinctly bent forward, bluish (?) in limb, wholly 
light yellow when dry, rarely glandular-pubescent on outside, glabrous 
inside, folds of lower lip densely hairy, lobes of lower lip ovate, 
subacute, slightly crenate; folds between them extending forward in 
the form of short teeth, as a result lower lip 5-lobed. Filaments 
subglabrous, puberulent at base; anthers subglabrous, uniformly hairy 
at base. Style subglabrous; stigrna yellow, bipartite. June to July. 

Host plant not determined.—Rubbly slopes.— Caucasus: Western 
Transcaucasia (eastern part), eastern Transcaucasia (western part), 
southern Transcaucasia (northern part). Endemic. Described from 
Borzhomi. Type in Leningrad. (Plate IV, fig. 4). 


15. O. serratocalyx G. Beck, Monogr. Orob. (1890) 104, tab. I, 
f. 15 and in Pflanzenr. IV, 261 (1930) 85.—Ic.: G. Beck, l.c. 

Biennial or perennial (?). Plant up to 25 cm tall, densely glandu- 
lar-hairy. Stem lilac, brownish whey dry, sometimes clavate at base, 
with relatively few ovate-lanceolate, about 1 cm long, scales. Inflores- 
cence elongated or shortly cylindrical, dense and multiflowered. Bract 
scales ovate, acuminate, often denticulate along margin, almost equal- 
ing calyx tube. Calyx densely glandular-hairy with distinct veins, teeth 
deltoid at base, subulate or narrowly acuminate, reaching middle of 
corolla tube and somewhat shorter than calyx tube, more or less serrate 
or dentate along margin. Corolla dilated from narrow part of tube 
toward limb, slightly bent forward, 20-24 mm long, limb deep bluish 
purple, usually reddish brown when dry, densely glandular-hairy on 
outside, upper lip deeply bilobate with triangularly subacute, crenate 
lobes, lower lip with orbicular or broadly ovate lobes, margin crenate 
or densely fimbriate. Filaments glabrous or with few short hairs at 
base and close to stigma; anthers sparsely hairy at base. Style densely 
glandular-puberulent. June to July. 

Host plant not known. Rubbly slopes. Not yet found in the USSR, 
but the collection of C. Koch was reported by Beck as found in the 
region of Turkey bordering Armenia. General distribution: Eastern 
Mediterranean (Syria), Armenia-Kurdistan. Described from Syria. Type 
in Vienna. 

Note. The systematic position of this species is still not clear. It 
probably belongs to the following subsection. 


Subsection 2. Pleioclada Novopokr. in Bot. mat. Gerb. Bot. 
inst. An SSSR, XIII (1950) 303, as section.—Stem branched, rarely 
simple, usually with few ovate or ovate-lanceolate scales. 
Predominantly annual or biennial. 


60 


233) 


16. O. hirtiflora (Reut.) Tzvel. comb. n.—O. aegyptiaca var. 
tricholoba G. Beck, Monogr. Orob. (1890) 103, tab. I, f. 14 and in 
Pflanzenr. IV, 261 (1930) 84.—? Phelipaea tricholoba Reut. in DC. 
Prodr. XI, 10 (excl. var. simplex).—P. hirtiflora Reut. |.c. 10; Boiss, 
Fl. or. IV, 497. 

Biennial or perennial (?). Plant 20-30 cm tall, glandular-pubes- 
cent, subglabrous below, however with grayish pubescence in inflores- 
cence, noticeable without magnification. Stem usually branched, yel- 
lowish, up to 8 mm thick in middle part, somewhat thicker at base, 
with few broadly lanceolate, up to 15 mm long, scales, often sinuate. 
Inflorescence short-cylindrical or cylindrical, lax, often dense in upper 
part, almost equaling or longer than remaining part of stem up to 20 
cm long. Bract scales lanceolate or broadly lanceolate, up to 18 mm 
long, usually slightly shorter than calyx, glandular-pubescent. 
Bracteoles linear-subulate, usually reaching middle of calyx teeth. 
Calyx 14-20 mm long, light brown, with relatively faintly prominent 
veins, glandular-pubescent, teeth deltoid at base, long subulate, usually 
longer than calyx tube (sometimes 2-3 times). Corolla 25-30 mm 
long, considerably dilated in limb, suberect, bluish (?), usually 
yellowish when dry, densely glandular-pubescent outside, glabrous 
inside except for sparsely hairy folds and lobes of limb, latter densely 
fimbriate and crenate along margin. Lobes of lower lip broadly ovate, 
sometimes subacute. Filaments shortly and sparsely pubescent only at 
base; anthers sparsely hairy at base. Style with isolated glandular- 
puberulent hairs. May to June. 

Host plant not precisely known; reported on species of Astragalus 
L. Rubbly slopes.— Caucasus: Southern Transcaucasia. General dis- 
tribution: Armenia-Kurdistan, Iran. Described from southern Iran. Type 
in Geneva. 

Note. Beck believes that Phelipaea tricholoba Reut. also belongs 
to this species, but it could hardly be true that the latter species either 
is a separate one, restricted to southern Iran, or is described from 
branched specimens of O. coelestis Boiss. et Reut., which is less 
probable. Both P. tricholoba and P. hirtiflora were described from the 
same collection number of Kotschy (Kotschy. Pl. Pers. austr. exs. No. 
408), containing a mixture of these species; hence, Beck could mistake 
P. hirtiflora as P. tricholoba. The separateness of O. hirtiflora (Reut.). 
Tzvel. cannot be doubted. 


17. O. aegyptiaca Pers. Synops. pl. II (1807) 181; G. Beck, 
Monogr. Orob. (1890) 100, tab. 1, f. 14 and in Pflanzenr. IV, 261 
(1930) 81, f. 12 (N) p.p.; Fedch. and Fier. Fl. 890, fig. 854; Grossh. 
Fl. Kavk. IV, 9.—O. indica Buchan. in Roxb. Fl. ind. III (1832) 27, 


6 


— 


56 


non Spreng. 1825; Ham. in Hook. FI. Brit. Ind. IV (1883) 326.—? O. 
longiflora Pers. |.c. 81; Schmalh. Fl. II, 294.—? Phelipaea longiflora 
C.A.M. Verzeichn. Pfl. cauc. (1831) 104; Reut. in DC. Prodr. XI, 5; 
Ldb. FI. Ross. III, 311; Boiss. Fl. or. IV, 495.—P. aegyptiaca Walp. 
Repert. III (1844-1845) 463; Reut. l.c. 9; Boiss. l.c. 499.—Ic.: G. 
Beck, I.c.; Fedch. and Fler. 1.c.—Exs.: GRF, No. 1427. 

Annual. Plant 15-50 cm tall, glandular-pubescent, sometimes 
subglabrous. Stem usually branched, yellowish, middle part up to 6— 
8 mm thick, slightly thickened at base; scales few, ovate-lanceolate, up 
to 15 mm long. Inflorescence cylindrical, lax, almost equaling re- 
maining part of stem or longer, up to 25 cm long. Bract scales ovate- 
lanceolate, 7-10 mm long, shorter than calyx, glandular-pubescent. 
Bracteoles linear-subulate, shorter than calyx. Calyx 8-14 mm long, 
usually extremely light colored, whitish, with less noticeable veins, 
glandular-pubescent, its teeth lanceolate-subulate, almost equaling calyx 
tube in length. Corolla 25-35 mm long, tubular-infundibuliform, con- 
siderably dilated in limb, suberect, bluish violet or deep blue, at base 
of tube lighter colored, almost whitish, sparsely glandular-pubescent 
outside, glabrous inside, except for sparsely hairy folds and lobes of 
limb, the latter with fimbriate and crenate margin; lobes of lower lip 
broadly ovate or orbicular, obtuse. Filaments hairy at base; anthers 
often densely hairy along sutures of theca as well as at base. Style 
glandular-puberulent. June to August (Plate IV, fig. 5). 

Predominantly parasitic on cultivated, occasionally on wild, plants 
belonging to various families of flowering plants (except for mono- 
cotyledonous plants, apparently erroneously indicated). Fields: cucur- 
bit fields, gardens; on wild plants usually along roadsides and around 
fields; rarely on rubbly slopes.—European USSR: Lower Volga, 
Crimea (reported by Beck, but absent in the herbaria of the Soviet 
Union). Caucasus: Southern Transcaucasia (Beck has reported it also 
in Dagestan, eastern Transcaucasia, Talysh). Soviet Central Asia: All 
regions besides northern part of Aralo-Caspian and Baltic regions. 
General distribution: eastern Mediterranean, Asia Minor, Armenia- 
Kurdistan, Iran, India-Himalayas, Dzhungaria-Kashgaria. Described 
from Egypt. Type in Leningrad. 

Note. Beck and A.A. Grossheim report for the Caucasus O. 
aegyptiaca var. aemula G. Beck [Monogr. Orob. (1890) 102], with 
smaller (20-26 mm long) flowers. In our opinion, however, plants of 
this variety are closer to O. mutelii F. Schultz, where consequently we 
place them. 

Economic importance. The plant is a serious pest of a whole 
range of agricultural crops, particularly cucurbits: melon, watermelon, 
cucumber, pumpkin. Within Solanaceae, it infects tomato, tobacco, 


62 


S| 


paprika, eggplant, rarely potato. Also observed on sunflower, cabbage, 
mustard, chinese bellflower, peanut, radish, rarely carrot, sesame, some 
ornamental plants. The list of wild host plants of Egyptian Orobanche 
is so large and diverse (according to the data of I.G. Beilin, D.T. 
Kabulov, V.V. Nikitin et al.) that it is not possible to cite them all 
here. For control measures, see Economic Importance of the genus. 


18. O. mutelii F. Schultz in Mutel, Fl. frang. II (1835) 353 and 
Atlas, tab. 43, f. 314; Suppl. tab. 2, f. 5 and in Flora, 67 and 168; G. 
Beck, Monogr. Orob. 95, tab. I, f. 13 and in Pflanzenr. IV, 261, 75; 
Fedch. and Fler. Fl. 890, fig. 893; Grossh. Fl. Kavk. IV, 9.—? 
Phelipaea macrantha C. Koch in Linnaea, XVII (1843) 290; Reut. in 
DC. Prodr. XI, II; Ldp. Fl. Ross. III, 312.—P. mutelii Reut. l.c. 8, 
p-p.—P. ramosa var. brevispicata Ldb. |.c. 313.—P. ramosa var. 
mutelii Boiss. Fl. or. IV (1879) 499, p.p.—Ic.: F. Schultz, I.c.; G. 
Beck, Monogr. Orob.; Fedch. and Fler. l.c. 

Annual or biennial. Plant 8-30 cm tall, glandular-pubescent. Stem 
branched or simple, yellowish or brownish, sometimes with red or 
blue color, middle part 2-5 mm thick, slightly thickened at base, with 
few ovate-lanceolate, up to 12 mm long, scales. Inflorescence 
cylindrical or ovate, lax or dense, usually exceeding or equaling, the 
remaining part of stem. Bract scales ovate or ovate-lanceolate, 6-10 
mm long, usually not reaching as far as upper teeth of calyx, glandular- 
pubescent. Bracteoles narrowly linear, somewhat shorter than calyx. 
Calyx 8-10 mm long, light brown, often wholly white, with slightly 
prominent veins, glandular-pubescent, its teeth lanceolate or subulate- 
lanceolate, equaling corolla tube or somewhat longer. Corolla usually 
18-20 mm, rarely up to 24 mm long, slightly curved forward, bluish- 
violet, lighter at base, glandular-pubescent outside; lobes of limb 
fimbriate and mildly crenate along margin, inside with sparse, short 
glandular hairs, orbicular or broadly ovate, obtuse; folds of lower lip 
densely hairy. Filaments subglabrous at base, with few short hairs; 
anthers hairy at base. Style subglabrous, or glandular-puberulent. May 
to August. 

Predominantly parasitic on various wild plants, rarely on culti- 
vated ones. Rubbly slopes, steppe, occasionally in fields and gardens.— 
European USSR: Crimea. Caucasus: All regions. General distribution: 
Atlantic Europe, Mediterranean, Balkans-Asia Minor, Armenia- 
Kurdistan, Iran. Described from France. Type in Paris. 

Note. A highly polymorphic species, differentiating into several 
physiological races showing slight morphological difference, which 
can be separated only in the presence of more abundant and 
thoroughly collected material. In addition to the most typical form 


63 


58 


(with a branched stem, relatively lax inflorescence and small flowers 
18-22 mm long), in the entire geographical range of the Caucasus two 
varieties are known to exist (probably, even distinct species): O. mutelii 
var. spissa (Rouy) G. Beck [Monogr. Orob. (1890) 96 = O. ramosa 
ssp. spissa Rouy in Bull. Soc. Bot. de France, LV (1908) 549] with a 
dense, shortly cylindrical inflorescence; simple stem and 18-22 mm- 
long flowers, parasitic only on wild plants, and O. mutelii var. interjecta 
G. Beck (Monogr. Orob. 98), somewhat intermediate between O. mutelii 
F. Schultz. and O. aegyptiaca Pers., with a lax inflorescence and larger 
flowers (up to 24 mm long), considerably dilated in the limb. In our 
opinion, specimens of O. aegyptiaca var. aemula G. Beck, reported 
from the Caucasus, belong to the last named of the foregoing varieties. 
Beck treated Phelipaea macrantha C. Koch (non Orobanche macrantha 
Dietr.), described from Armenia by C. Koch, as a synonym of O. 
mutelii var. spissa (Rouy) G. Beck; however, probably it would be 
more correct to place this species [P. macrantha] in the synonymy of 
O. cilicica G. Beck. Apparently, reports of O. mutelii P. Schultz for 
Soviet Central Asia are erroneous. 

Economic importance. Sometimes damages agricultural crops, 
parasitizing tobacco, tomato, mustard, vetch, clover, peavine, and some 
ornamental plants. Among ornamental plants, the most vulnerable are 
primarily species of the Compositae, Leguminosae, Umbelliferae, 
Cruciferae, and Solanaceae, as well as of many other families. For 
preventive measures, see Economic Importance of the genus. 


19. O. brassicae Novopokr. in Izv. Donsk. inst. sel’sk. khoz. 1 
melior. IX (1929) 47, 54, 58; Gonch. in Sorn. rast. SSSR, IV, 135 and 
in Fl. Yugo-Vost. VI, 232; Grossh. Opred. rast. Kavk. 353.—O. mutelii 
ssp. brassicae Novopokr. |.c. VIII (1928) 52, fig. 1.—Ic.: Novopokr. 
l.c.; Beilin, Zarazikhi i bor’ba s nimi, fig. 15 (as O. mutelii F. 
Schultz).—Exs.: GRF, No. 3489. 

Annual. Plant 15-30 cm tall, glandular-hairy. Stem branched, rarely 
simple, bluish gray, when dry brownish, middle part 3-12 mm thick, 
base usually tuberously thickened, scales few, ovate-lanceolate but 
usually long acuminate, 10-12 mm long. Inflorescence cylindrical, 
rarely short cylindrical, lax or fairly dense, usually equaling or ex- 
ceeding the remaining part of stem. Bract scales ovate-lanceolate, acute, 
usually not reaching tips of calyx teeth, 7-10 mm long, glandular- 
hairy. Bracteoles linear-subulate, shorter than calyx. Calyx 9-10 mm 
long, light brown, often wholly whitish with slightly prominent veins, 
glandular-pubescent, teeth triangular at base to lanceolate-subulate, 
somewhat exceeding calyx tube. Corolla 18—24 mm long, slightly curved 
forward, with dark violet limb and paler tube, whitish at base, glandu- 


64 


59 


lar-pubescent outside; lobes of limb with fimbriate and mildly crenate 
margin, sparsely glandular-pubescent inside, orbicular or broadly ovate, 
obtuse; folds of lower lip densely hairy. Filaments subglabrous, with 
few short hairs at base; anthers glabrous, rarely with several hairs at 
base. Style subglabrous. August to September. 

Parasitic on cabbage (Brassica oleracea L.), tomato (Lycopersicum 
esculentum Mill.), rape (Brassica napus L.), mustard (Sinapus alba 
L.), and occasionally probably also on wild plans. Fields, gardens. — 
European USSR: Bessarabia, Black Sea Region, Lower Don, Crimea. 
General distribution: Probably occurs in Romania and Bulgaria. De- 
scribed from Rostov Region. Type in Novorossisk, topotype in 
Leningrad. 

Note. Evidently, O. mutelii F. Schultz appeared relatively recently 
in agriculture and is a more northern physiological race. Morphologi- 
cal differences between these species are insignificant. 

Economic importance. Currently seen only in several regions, 
but sometimes abundant, being (in this case) harmful to tomato and 
cabbage. 


20. O. hohenackeri (Reut.) Tzvel. comb. n.—O. mutelii auct. 
p.p.—Phelipaea hohenackeri Reut. in DC. Prodr. XI (1847) 10.—P. 
ramosa var. crassipes Ldb. Fl. Ross. III, 313. 

Annual or biennial. Plant up to 30 cm tall, glandular-pubescent. 
Stem with inflorescence usually branched, rarely simple, yellowish or 
brownish, middle part up to 8—9 mm thick, base slightly thicker; scales 
relatively few, broadly lanceolate, up to 15 mm long. Inflorescence 
cylindrical or oblong, fairly dense, lax in lower part, longer than or 
equaling remaining part of stem. Bract scales lanceolate, usually not 
reaching tip of calyx teeth, 10-15 mm long. Bracteoles linear-subu- 
late. Calyx 12-14 mm long, light brown, often wholly white, with 
slightly noticeable veins, teeth of calyx lanceolate at base to subulate, 
1%4-2% times as long as calyx tube, deflexed to a side in fruit, deeply 
incised on the side of inflorescence axis, reaching nearly to calyx 
base. Corolla 18-20 mm long, in limb blue or violet (?), uniformly 
brownish yellow when dry, subglabrous outside with sparse, extremely 
short glandular hairs; lobes of lower lip crenate, broadly ovate, 
sometimes subacute; folds hairy. Anthers sparsely hairy; filaments 
sparsely pubescent at base, sometimes subglabrous. Style glandular- 
puberulent. May to June. 

Host plant unknown. Rubbly slopes.—Caucasus: Western 
Transcaucasia(?), Talysh. General distribution: Armenia-Kurdistan, 
Iran. Described from Talysh (Zuvand). Type in Paris(?), isotype in 
Leningrad. 


65 


60 


Note. Besides several authentic specimens of O. hohenackeri from 
Zuvand, the Herbarium of the Botanical Institute of the Academy of 
Sciences of the USSR also possesses one specimen of this species 
collected by Eichwald, probably in western Transcaucasia. 


21. O. brachypoda Novopokr. in Bot. mat. Gerb. Bot. inst. AN 
SSSR, XIII (1950) 303. 

Biennial or annual. Plant up to 20 cm tall, glandular-pubescent. 
Stem branched in inflorescence, rarely simple, middle part 3-7 mm 
thick, base usually thickened like club, yellow, light brown when dry; 
scales several, caducous. Inflorescence cylindrical or elongated with a 
few short branches at base, dense in upper part, lax below, with 
distantly placed flowers, considerably longer than highly shortened 
stem, usually starting from the ground itself. Bract scales broadly 
lanceolate, not reaching tip of calyx teeth, 5-10 mm long. Bracteoles 
linear-subulate. Calyx 7-9 mm long, light brown, with bluish tinge, 
its teeth narrowly lanceolate, almost equaling calyx tube. Incision of 
calyx from side of inflorescence axis 2-3 mm, not reaching its base. 
Corolla 20-22 mm long, limb deep blue or bluish, densely glandular- 
pubescent outside, lobes of lower lip broadly ovate, sometimes 
subacute; folds sparsely hairy. Filaments sparsely pubescent at base; 
anthers densely hairy at base and often up to sutures of thecae. Style 
glandular-puberulent. June to July. 

Host plant not determined.—Rubbly slopes.— Soviet Central Asia: 
Pamiro-Alai. Endemic. Described from Turkestan Range. Type in 
Leningrad. 


22. O. oxyloba (Reut.) G. Beck in L. Koch, Entwickl.-Gesch. der 
Orob. (1887) 209; Monogr. Orob. (1890) 108, p.p., tab. I, f. 18 (2); 
in Pflanzenr. IV, 261 (1930) 89, p.p.; Grossh. Fl. Kavk. IV, 9, p.p.— 
Phelipaea oxyloba Reuter in DC. Prodr. XI (1847) 9; Boiss. Fl. or. IV, 
497.—Ic.: G. Beck, l.c. 

Annual or biennial. Plant up to 15—20 cm tall, glandular-pubes- 
cent. Stem simple or branched, yellowish or light brown, with few 
ovate-lanceolate scales. Inflorescence short cylindrical or ovate, lax 
and relatively few-flowered, usually exceeding or equaling remaining 
part of stem. Bract scales ovate-lanceolate, 6-9 mm long. Bracteoles 
linear-subulate. Calyx 7-9 mm long, light brown, often whitish, with 
less prominent viens, calyx teeth lanceolate-subulate, almost equaling 
calyx tube. Corolla 17-20 mm long, in limb light-violet (?), usually 
yellowish when dry, sparsely glandular-pubescent outside, lobes of 
lower lip crenate, broadly ovate, acuminate; folds hairy. Filaments 


6 


nN 


61 


nearly glabrous; anthers glabrous or sparsely hairy at base. Style 
sparsely glandular-puberulent. May to July. 

Host plant not determined. Rubbly slopes, sometimes in forests.— 
European USSR: Crimea (southern part). Caucasus: Western 
Transcaucasia, Talysh. General distribution: Asia Minor, Armenia- 
Kurdistan, Iran. Described from Turkey. Type in Paris (?), isotype in 
Leningrad. 


23. O. orientalis G. Beck, Monogr. Orob. (1890) 110, tab. 1, f. 
19 and in Pflanzenr. IV, 261 (1930) 90.—Ic.: G. Beck, l.c. 

Biennial or perennial. Plant up to 30—40 cm tall, densely glandular- 
pubescent. Stem branched, rarely simple, brownish yellow, with 
relatively few or fairly numerous ovate or ovate-lanceolate scales. 
Inflorescence cylindrical lax in lower part, with distant flowers, dense 
in upper part, multiflowered, pointed at tip, usually longer than highly 
shortened part of stem. Bract scales ovate-lanceolate, 6-11 mm long. 
Bracteoles linear-subulate. Calyx 7-12 mm long, light brown, teeth 
lanceolate-subulate, usually longer than calyx tube, rarely almost as 
long, somewhat deflexed in fruit. Corolla 14-19 mm long, pale blue- 
pinkish, whitish at base, glandular-pubescent outside, lobes of lower 
lip ovate, subacute, nearly entire or finely crenate; folds of lower lip 
usually hairy. Filaments glabrous or subglabrous; anthers densely hairy 
at base and along sutures of thecae. 

Parasitic on the Amygdalus L. species; in the Caucasus on A. 
fenzliana (Fritsch) Lipsky. Rubbly slopes, scrub of lower and central 
mountain belts.— Caucasus: Eastern Transcaucasia (southeastern part). 
Soviet Central Asia; Pamiro-Alai. General distribution: Iran- 
Afghanistan, India-Himalayas. Described from Afghanistan. Type in 
London, isotype in Leningrad. 

Note. Specimens collected in the Caucasus in the regions of Mount 
Megra (Armenian SSR) and the village of Zangelan (Nagorno-Karabakh 
Autonomous Region) are completely similar to plants of O. orientalis 
G. Beck from Tadzhikistan, differing only in somewhat more narrow 
and more acute corolla lobes. However, they are also similar to an- 
other related species—O. schultzii Mutel (Fl. franc. (1835) 352 and 
429), specimens of which were collected in the Near East (identified 
by Beck) and are also represented at the Herbarium of the Botanical 
Institute of the Academy of Sciences of the USSR. Having in mind 
that this latter species has been described from Algeria and reported 
as a parasite of legumes, we prefer to treat this Caucasian plant as O. 
orientalis G. Beck. The next species, evidently, also represents a 
morphologically weakly differentiated (the whole plant less pubescent) 
physiological race of O. orientalis G. Beck. 


69 


62 


24. O. sogdiana Novopokr. in Bot. mat. Gerb. Bot. inst. AN SSSR, 
XIII (1950) 306. 

Biennial or perennial (?). Plant 10-30 cm tall, glandular-pubes- 
cent. Stem simple or branched, yellowish brown when dry, middle 
part 2-4 mm thick; scales relatively few, ovate or ovate-lanceolate, up 
to 10 mm long. Inflorescence cylindrical or oval, dense in upper part, 
fairly lax in lower part, multiflowered, tip usually rounded, equaling 
or longer than remaining part of stem. Bract scales ovate-lanceolate. 
Bracteoles linear-subulate. Calyx 8-10 mm long, light brown, calyx 
teeth lanceolate-subulate, equaling calyx tube or somewhat longer. 
Corolla 15-18 mm long, pale blue, whitish at base, sparsely pubescent 
outside, often subglabrous; lobes of lower lip broadly oval, subacute, 
almost entire; folds hairy. Filaments glabrous; anthers hairy at base. 
Style glabrous. May to July (Plate IV, fig. 6). 

Parasitic on Amygdalus bucharica Korsh. Rubbly slopes, scrub.— 
Soviet Central Asia: Pamiro-Alai. Endemic. Described from Hissar 
Mountains. Type in Leningrad. 


25. O. dalmatica (G. Beck) Tzvel. comb. n.— O. oxyloba var. 
dalmatica G. Beck, Monogr. Orob. (1890) 109, tab. 1, f. 18(1) and in 
Pflanzenr. IV, 261 (1930) 89; Grossh. Fl. Kavk. IV, 9.—Ic.: G. Beck, 
ke 
Biennial or annual (?). Plant up to 25 cm tall, glandular-pubescent, 
sometimes subglabrous below. Stem simple or branched in inflores- 
cence, reddish, 2-5 mm thick in middle part, with few lanceolate, 7— 
9 mm long, scales. Inflorescence short cylindrical or lanceolate, dense 
and multiflowered, subacute, almost equaling remaining part of stem, 
with short lateral branches at base. Bract scales broadly lanceolate, 6— 
8 mm long. Bracteoles linear-subulate. Calyx 7-9 mm long, light brown 
with darker, but weakly prominent veins; teeth triangular at base, subu- 
late, equaling calyx tube or somewhat shorter. Corolla 16-18 mm long, 
with bluish or violet (?) limb, usually yellow when dry, sparsely glan- 
dular-puberulent outside; lobes of lower lip oval, subacute, margin 
almost entire; folds usually hairy. Filaments sparsely pubescent at base; 
anthers pubescent at base. Style subglabrous. May to July. 

Host plant not determined. Rubbly slopes. — Caucasus: Ciscaucasia, 
western Transcaucasia (northern part). General distribution: Balkans- 
Asia Minor. Described from Yugoslavia. Type in Vienna. 

Note. We consider this species closer to O. schultzii Mutel. and to 
several forms of O. nana Noé than to O. oxyloba (Reut.) G. Beck. Of 
the Caucasian specimens of this species identified by Beck, the one 
from the vicinity of Anapa differs somewhat from two others from the 


Plate V. 

67 1. Orobanche amurensis (G. Beck) Kom., flower.—2. O. amoena C.A.M., 
flower. —3. O. cumana Wallr., flower.—4. O. camptolepis Boiss. and Reut., 
general appearance of plant, flower.—5. O. sordida C.A.M., flower.—6. O. 
gamosepala Reut., flower.—7. O. lutea Baumg., flower—8. O. alba Steph., 
flower.—9. O. raddeana G. Beck, flower.—10. O. hederae Duby, flower. — 

11. O. inulae Novopokr. and Abram., general appearance of plant, flower. 


70 


64 


vicinity of Pyatigorsk and Groznyi. The latter specimens are imperfect, 
and it is doubtful whether they belong to this species. 


26. O. nana Noé in Rchb. Herb. norm. No. 1352; G. Beck, Monogr. 
Orob. (1890) 91, tab. 1, f. 11 and in Pflanzenr. IV, 261 (1930) 70; 
Fedch. and Fler. Fl. 889, fig. 852, Grossh. Fl. Kavk. IV, 9.—Phelipaea 
muteli var. nana Reuter in DC. Prodr. XI (1847) 9.—P. ramosa var. 
nana Boiss. Fl. or. IV (1879) 499.—Ic.: G. Beck, l.c.; Fedch. and 
Flermalic: 

Annual or biennial. Plant 15—20 cm tall, glandular-pubescent. Stem 
simple, rarely branched, yellowish, often clavate at base, with few 
ovate-lanceolate, about 8 mm long, scales. Inflorescence oval or short 
cylindrical, dense or lax, often few-flowered, equaling, shorter or 
slightly longer than remaining part of stem. Bract scales ovate- 
lanceolate, up to 8 mm long. Bracteoles linear-subulate. Calyx 5-8 
mm long, usually whitish, with weak veins, calyx teeth deltoid, rarely 
lanceolate at base, gradually subulate, equaling calyx tube or somewhat 
longer. Corolla 10-17 mm long, blue or violet in limb, glandular- 
pubescent outside; lobes of lower lip orbicular or broadly oval, obtuse; 
folds densely hairy. Filaments sparsely pubescent at base, subglabrous; 
anthers glabrous or sparsely hairy at base. Style sparsely glandular- 
puberulent under stigma. May to July. 

Parasitic on various wild plants, usually on annuals (reported on 
species of genera Torilis Adans., Trigonella L., Lamium L., Veronica 
L., and many others). Rubbly slopes, scrub.— European USSR: Lower 
Don (southern part), Crimea (southern part). Caucasus: All regions. 
Soviet Central Asia: Pamiro-Alai (western part). General distribution: 
Mediterranean, Balkans-Asia Minor, Armenia-Kurdistan, Iran. 
Described from Yugoslavia. Type in Leipzig, isotype in Leningrad. 

Note. The differences between this and the following species are 
not always clearly expressed. Evidently O. nana Noé is ancestral to 
the relatively younger species O. ramosa L., which has adapted to 
parasitizing cultivated plants. 


27. O. ramosa L. Sp. pl. (1753) 633; M.B. Fl. taur.-cauc. II (1808) 
83; G. Beck, Monogr. Orob. 87, tab. 1, f. 10 and in Pflanzenr. IV, 
261, 66, f. 12 (A); Schmalh. FI. II, 294; Fedch. and Fler. Fl. 889, fig. 
850 and 851; Grossh. Fl. Kavk. IV, 8.—Phelipaea ramosa C.A.M. 
Verzeichn. Pfl. Cauc. (1831) 104; Reut. in DC. Prodr. XI, 8; Ldb. Fl. 
Ross. III, 313; p.p.; Boiss. Fl. or IV, 498, p.p.—Ic.: Rcehb. Ic. Fl. 
Germ. XX, tab 152; G. Beck, 1.c.; Hegi. Ill. Fl. VI, f. 80; Fedch. and 
Fler., l.c.; Maltsev, Atlas vazhn. sorn. rast. SSSR, II (1939), plate 40 
and fig. 90. Exs.: GRF, No. 378; Fl. pol. exs. No. 857. 


71 


65 


Annual. Plant up to 25(40) cm tall, glandular-pubescent. Stem 
branched, rarely simple, yellowish, base usually not thickened, with 
few ovate scales. Inflorescence cylindrical or lanceolate, fairly lax, 
longer than or equaling remaining part of stem. Bract scales ovate- 
lanceolate, 5-8 mm long. Bracteoles linear-subulate. Calyx 5-8 mm 
long, usually whitish, with weak veins, calyx teeth narrowly lanceolate 
or lanceolate-subulate, equaling calyx tube or shorter, rarely somewhat 
longer. Corolla 10-17 mm long, pale violet, light yellow at base of 
tube, glandular-pubescent outside; lobes of lower lip orbicular or 
broadly oval, obtuse; folds densely hairy. Filaments sparsely pubescent 
at base, subglabrous; anthers glabrous or sparsely hairy at base. Style 
sparsely puberulent under stigma. July to August (Plate IV, fig. 7). 

Parasitic on various cultivated and wild plants, predominantly 
annuals. Fields, gardens; on nearby wild plants also, roadsides, old 
fields.— European USSR: Upper Dniester, Upper Dnieper, Middle 
Dnieper, Volga-Don, Bessarabia, Black Sea Region, Lower Don, 
Crimea.— Caucasus: All regions. General distribution: Central and 
Atlantic Europe, Mediterranean, Armenia-Kurdistan, Iran; introduced 
to North America. Described from Europe. Type in London. 

Economic importance. The species is harmful to certain cultivated 
plants, especially, hemp, tobacco, tomato. Also observed on musk- 
melon, gourd, carrot, coriander, sunflower, Russian dandelion, mustard, 
cabbage, potato, beans, perilla, peanut. 


Section 2. OsPpROLEON Wallroth. Sched. crit. I (1822) 307; G. Beck, 
Monogr. Orob. 133 and in Pflanzenr. IV, 261, 113.—Orobanche auct. 
non L.: C.A.M. in Ldb. FI. alt. II (1830) 450; Reut. in DC. Prodr. XI, 
15; Ldb. FI. Ross. III, 315; Boiss. Fl. or. 1V, 502.—Calyx incised into 
2 lateral entire or doubly serrated segments up to base, or nearly so, 
very rarely gamosepalous, 4—5 toothed. Bracteoles absent. All flowers 
in inflorescence sessile. Stem always simple. 


Subsection 1. Inflatae G. Beck in Halacsy u. Braun, Nachr. 
zur Fl. Nieder-Oesterr. (1822) 124; Monogr. Orob. (1890) 136 and in 
Pflanzenr. IV, 261 (1930) 115.—Corolla with elongated tube usually 
more or less inflated below place of insertion of stamens, thereafter 
narrowed and then again spreading out towards limb. Stamens inserted 
in corolla tube, one-third from base or above, sometimes almost at 
middle. of tube. 

Note. Beck proposed dividing this subsection into two groups: 
Coerulescentes G. Beck and Amoenae G. Beck. We consider a division 
into series more natural. 


Vz 


66 


Series 1. Coerulescentes G. Beck, Monogr. Orob. (1890) 136, p.p. 
(sub grege).—Araneosae Novopokr. in Bot. mat. Gerb. Bot. inst. AN 
SSSR (1950) 311, p.p.—Corolla tube distinctly curved forward, with 
constriction below the point of insertion of stamens. Stamens inserted 
somewhat below its middle; anthers glabrous. Plant white lanate or 
glabrous in inflorescence. Parasitic on species of sagebrush. 


28. O. coerulescens Steph. in Willd. Sp. pl. II (1800) 349; M.B. 
Fl. taur.-cauc. II, 82 and III, 417; Reut. in DC. Prodr. XI, 34; Ldb. FI. 
Ross. III, 322; G. Beck, Monogr. Orob. 137, tab. II, f. 81 and in 
Pflanzenr. IV, 161, 118, f. 13 (A); Schmalh. Fl. II, 297; Kom. FI. 
Manchzh, III, 467; Fedch. and Fler. Fl. 891, fig. 859; Kom. and Allis. 
Opred. rast. Dalnevost. Kr. II, 936, plate 279; Kryl. Fl. Zap. Sib. X, 
2539.—O. ammophila C.A.M. in Ldb. FI. alt. II (1830) 454; Reut. l.c. 
36.—Ic.: Ldb. Ic. pl. fl. Ross. imp. alt. IV (1833) tab. 389; G. Beck, 
l.c.; Fedch. and Fler. I.c.; Hegi, Ill. Fl. VI, f. 82 (a—-c); Kom. and Allis. 
l.c. 

Perennial. Plant up to 40 cm tall, more or less white arachnoid- 
lanate. Stem yellowish, middle part 2.5-10 mm thick, usually claviform 
at base, scales sessile, lanceolate or ovate-lanceolate, up to 20 mm 
long. Inflorescence cylindrical or oval, dense, rarely lax. Bract scales 
ovate or ovate-lanceolate, upper part often extended into, up to 20 
mm long, linear tongue. Calyx segments free, 8-13 mm long, entire 
or split into 2 narrow teeth at most up to middle. Corolla 15—20 mm 
long, blue or violet, usually preserving its color in part when dry, 
tubular, distinctly curved forward, arachnoid-villous outside, glabrous 
inside, folds of lower lip sparsely hairy. Filaments glabrous or sparsely 
hairy at base, inserted at distance of 6-7 mm from the base of corolla 
tube, somewhat below its middle; anthers glabrous (extremely rarely 
hairy at base). Style sparsely glandular-puberulent under stigma. May 
to July. 

Parasitic on species of sagebrush (Artemisia marschalliana Spreng., 
A. commutata Bess., A. frigida Willd., A. dracunculus L., A. glauca 
Pall., A. latifolia Ldb., etc.). Steppe, rubbly slopes, sandy areas.— 
European USSR: Volga-Kama (southern part), Upper Dniester, Middle 
Dniester, Volga-Don, Volga Region, Bessarabia, Black Sea Region, 
Lower Don, Lower Volga. Caucasus: Ciscaucasia, Dagestan. Western 
Siberia: Upper Tobol, Irtysh, Altai. Eastern Siberia: Angara-Sayan, 
Dauria, Lena-Kolyma (southern part). Soviet Far East: Zeya-Bureya, 
Uda River area, Ussuri. Soviet Central Asia: Aralo-Caspian, Balkhash 
Region, Dzhungaria-Tarbagatai. General distribution: Central Europe, 
Balkans, Dzhungaria-Kashgaria, Mongolia, Japan, China. Described 
from the area of Lake Inder. Type in Leningrad. 


73 


67 


29. O. korshinskyi Novopokr. in Bot. mat. Gerb. Bot. inst. AN 
SSSR, XIII (1950) 311.—P. coerulescens var. albiflora O. Kuntze in 
Tr. SPb., bot. sada, X, 225; Korsh. Tentam. fl. Ross. or. 327 (pro 
forma). 

Perennial. Plant 15-35 cm tall, more or less white arachnoid- 
lanate. Stem light yellow, middle part 2.5-5 mm thick, usually 
claviform at base; scales sessile-lanceolate or ovate-lanceolate, 10-17 
mm long. Inflorescence oval or shortly cylindrical, fairly dense, rarely 
lax. Bract scales ovate or ovate-lanceolate. Calyx segments free, 8-12 
mm long, entire or bidentate, with narrowly lanceolate teeth. Corolla 
14-18 mm long, light yellow, tubular, curved forward (not as much as 
that compared with O. coerulescens Steph.), more or less whitish, 
arachnoid-villous outside. Filaments almost glabrous, inserted at a 
distance of about 6 mm from base of corolla tube; anthers glabrous. 
Style subglabrous, glandular-puberulent under stigma. June to July. 

Parasitic on species of Artemisia (A. sericea Web., A. latifolia 
Ldb., A. laciniata Willd., A. marschalliana Spreng., A. tanacetifolia 
Denetey: 

Steppe, rubbly slopes, sands.— European USSR: Volga-Kama 
(southern part), Volga Region, Lower Volga. Western Siberia: Upper 
Tobol, Irtysh, Altai. Eastern Siberia: Angara-Sayan, Dauria. Soviet 
Far East: Zeya-Bureya, Ussuri. General distribution: Mongolia, Japan- 
China. Described from Southern Urals (Orenburg Region). Type in 
Leningrad. 

Note. Possibly a color variant of the foregoing species. 


30. O. glaucantha Trautv. in Tr. SPb., bot. sada, I (1871) 26. 

Perennial or biennial (?). Plant up to 35 cm tall, glabrous. Stem 
yellowish, sometimes with reddish tinge, middle part up to 6 mm 
thick, slightly thicker at base, with fairly numerous lanceolate, up to 
18 mm long, scales. Inflorescence oblong or short cylindrical, fairly 
dense, usually 24% times as long as remaining part of stem. Bract 
scales lanceolate, up to 15 mm long. Calyx segments free, 8-12 mm 
long, entire or bidentate at most up to middle. Corolla 15-18 mm 
long, blue, usually preserving its color partially when dry, tubular, 
distinctly curved forward, glabrous. Filaments glabrous, inserted at 
distance of 6-8 mm from base of corolla tube; anthers and style 
glabrous. June to July. 

Host plant not determined; probably on species of Artemisia 
Rubbly slopes.— Eastern Siberia: Angara-Sayan (upper reaches of 
Angara River). Endemic. Described from Irkutsk Region. Type in 
Leningrad. 


74 


68 


Note. Despite the complete absence of pubescence, this species is 
extremely close in every other respect to O. coerulescens Steph., as 
though representing its glabrous variant. However the presence of 
several specimens of this species from the same region, in the Herbarium 
of the Botanical Institute of the Academy of Sciences of the USSR is 
evidence of the fact that absence of pubescence is not accidental in 
this case. Since Beck did not come across specimens of O. glaucantha 
Trautv., he erroneously referred this species to O. cumana Wallr. 


Series 2. Pycnostachyae Tzvel. Corolla tube narrow and long, 
distinctly curved forward at place of attachment of stamens, almost 
erect toward limb with very slight constriction. Stamens inserted 
in corolla tube at a point slightly above one third from base; 
anthers densely hairy. Plant glandular-pubescent. Parasitic on Artemisia 
species. 


31. O. pycnostachya Hance in Journ. Linn. Soc. XIII (1873) 84; 
G. Beck, Monogr. Orob. 140, tab. II, f. 32 and in Pflanzenr. IV, 261, 
116; Kom. Fl. Manchzh. III, 468; Kom. and Alis. Opred. rast. Dalnevost. 
kr. II, 935.—O. macrolepis Turcz. Catal. baical. No. 878; Ldb. FI. 
Ross. III, 323 (nomen); Turcz. in Bull. Soc. Nat. Mosc. XXIV, 356, 
non Coss. 1847. 

Perennial or biennial (?). Plant up to 45 cm tall, glandular- 
pubescent. Stem brownish yellow, middle part 2-5 mm thick, slightly 
thickened at base; scales sessile, lanceolate, up to 18 mm long. 
Inflorescence cylindrical or oblong, fairly dense and multiflowered, 
usually shorter than remaining part of stem. Bract scales broadly 
lanceolate, up to 22 mm long. Calyx segments free, 10-15 mm long, 
usually bidentate, with narrowly lanceolate teeth, rarely entire. Corolla 
17-25 mm long, with long and narrow tube, distinctly curved forward, 
slightly patent in limb, light yellow, sometimes with faded bluish color 
in limb, glandular-pubescent outside, inside with sparse glandular hairs 
on lobes and folds of lower lip. Filaments at base with a short, sparse 
hairs, inserted at a distance, of 5.7 mm from base of corolla tube; 
anthers densely hairy at base and along sutures of thecae. Style 
glandular-puberulent. June to July. 

Parasitic on wormwood (Artemisia sacrorum Ldb., A. commutata 
Bess., etc.). Rubbly slopes, scrub, dry meadows.— Eastern Siberia: 
Dauria. Soviet Far East: Zeya-Bureya, Ussuri. General distribution: 
Japan, China. Described from China. Type in London. 


75 


69 


32. O. amurensis (G. Beck) Kom. Fl. Manchzh. III (1907) 469; 
Kom. and Alis. Opred. rast. Dalnevost. kr. II, 936.—O. pycnostachya 
var. amurensis G. Beck, Monogr. Orob. (1890) 141 and in Pflanzenr. 
IV, 261 (1930) 118.—O. ussuriensis Novopokr. in Bot. mat. Gerb. 
Bot. inst. AN SSSR, XII (1950) 274. fig. 1.—Ic.: Novopokr. l.c. 

Perennial or annual (?). Plant up to 45 cm tall, glandular-pubescent. 
Stem brownish yellow, middle part 2-6 mm thick, slightly thicker at 
base; scales sessile, lanceolate, up to 20 mm long. Inflorescence 
cylindrical or oblong, usually lax, rarely thick, multiflowered, equaling 
longer or shorter than remaining part of stem. Bract scales broadly 
lanceolate, but often reflexed at apex and narrower with linear tip, 
15-20 mm long. Calyx segments free, 8-18 mm long, usually incised 
into 2 narrowly lanceolate or lanceolate-subulate teeth up to middle or 
even deeper, rarely entire. Corolla 18—25 mm long, with long, narrow 
tube, distinctly (but less marked as compared to O. pycnostachya 
Hance) curved forward, slightly broadened in limb, blue or violet, 
often preserves color, even when dry, glandular-pubescent outside. 
Filaments sparsely pubescent at base; anthers densely hairy at base 
and usually along sutures of thecae. Style subglabrous. June to July 
(Plate V, fig. 1). 

Parasitic on species of wormwood (Artemisia sacrorum Ldb., A. 
laciniata Willd., A. commutata Bess., etc.). Rubbly slopes, shingles, 
dry meadows.—Soviet Far East: Zeya-Bureya, Ussuri. General 
distribution: Japan, China. Described from Primorye Region. Type in 
Leningrad. 

Note. O. ussuriensis Novopokr., having slightly more sparse 
pubescence over entire plant, is in no other way different from this 
species and, in our opinion, cannot be regarded even as a distinct 
variety. 


Series 3. Amoenae G. Beck, Monogr. Orob. (1890) 133, 148, p.p. 
(sub tribu) and in Pflanzenr. IV, 261 (1930) 131, p.p. (sub grege).— 
Corolla tube slightly bent, often suberect, slightly constricted above 
the place of insertion of stamens, limb of corolla, blue, violet, or deep 
blue. Stamens usually inserted in lower third of corolla tube. Anthers 
sparsely hairy. Plant glandular-pubescent. Parasitic on composites. 


33. O. amoena C.A.M. in Ldb. FI. alt. II (1830) 457; Reut. in 
DC. Prodr. XI, 31; Ldb. Fl. Ross. III, 320; G. Beck, Monogr. Orob. 
148, tab. II, f. 37 and in Pflanzenr. IV, 261 (1930) 131, f. 13 (F-H); 
Gonch. in Fl. Yogo-Vost. VI, 238; Kryl. Fl. Zap. Sib. X, 2542.—0O. 
colossea Novopokr. in Bot. mat. Gerb. Bot. inst. AN SSSR, XII (1950) 


70 


276.—O. comosula Novopokr., |.c. 280.—Ic.: Ldb. Ic. pl. fl. Ross. 
imp. alt. IV (1833), tab. 385; G. Beck, Monogr. Orob. |.c.—Exs.: Kar 
and Kir. exs. ann. 1840, No. 382 and 385. 

Perennial or biennial. Plant up to 40 cm tall, glandular-pubescent, 
often with violet or reddish tint. Stem 2—8 mm thick in middle, usually 
slightly thicker at base; scales sessile, ovate-lanceolate or lanceolate, 
12-18 mm long. Inflorescence oval or short cylindrical, usually fairly 
dense, rarely lax, often shorter than, or equaling, remaining part of 
stem. Bract scales ovate-lanceolate, 7-18 mm long. Calyx segments 7— 
15 mm long, light brown, often whitish, entire or bidentate; teeth 
narrowly lanceolate, often subulate at tip. Corolla 20-40 mm long, 
tubular, from constricted part of tube to collar, gradually but 
considerably broadened, blue or violet (sometimes only in limb), 
preserving color partially when dry, sparsely glandular-puberulent or 
subglabrous outside. Filaments pubescent at base, inserted, at distance 
of 6-11 mm from base of corolla tube; anthers pubescent along sutures 
of thecae. Style subglabrous. June to July (Plate V, fig. 2). 

Parasitic on wormwood species (Artemisia glanduligera Krasch., 
A. porrecta Krasch., A. lessingiana Bess. etc., usually on section 
Seriphidium Bess.). Steppe, rubbly slopes, alkali soils — European 
USSR: Trans- Volga, Lower Volga (northeastern part). Western Siberia: 
Upper Tobol, Irtysh, Altai. Soviet Central Asia: All regions. General 
distribution: Iran-Afghanistan (northeastern part), Dzhungaria- 
Kashgaria, Mongolia (western part). Described from Altai. Type in 
Leningrad. 

Note. A polymorphic species, varying greatly in the size and color 
of the corolla, extent of cutting of the calyx segments, form of the 
inflorescence and other characters. Beck distinguishes seven forms of 
this species. The specimens with the largest corolla (35-40 mm long), 
found in the mountains of Soviet Central Asia, together with the typical 
form, are referred to f. colossea G. Beck (= O. colossea Novopokt.). 
O. comosula Novopokr. evidently was described from young flowering 
specimens which, in our opinion, do not differ essentially from O. 
amoena C.A.M. 


34. O. ariana Gontsch. in FI. i sistem; vyssh. rast. I (1933) 173. 
Perennial or biennial (?). Plant up to 25 cm tall, glandular- 
pubescent. Stem up to 9 mm thick in middle part, usually slightly 
thicker at base, with few sometimes obtuse, up to 11 mm long, scales. 
Inflorescence extremely dense, multiflowered, rounded at tip, equaling 
remaining part of stem or somewhat shorter. Bract scales lanceolate, 
about 15 mm long, lower ovate and about 10 mm long. Calyx segments 
10-11 mm long, incised up to middle or deeper into two long and thin 

76 acuminate teeth. Corolla 24-26 mm long, whitish, pale violet in limb, 


71 


campanulate-tubular, collar infundibular, easily bent forward, sparsely 
glandular-puberulent outside. Filaments hairy at base, inserted at height 
of 7-8 mm from base of corolla tube; anthers sparsely hairy along 
sutures of thecae. Style sparsely glandular-puberulent. June to July. 

Parasitic, evidently, on species of Artemisia L. Rubbly slopes.— 
Soviet Central Asia: mountainous Turkmenia. General distribution: 
probably Iran. Described from Kopet-Dag (near Akar-Cheshme). Type 
in Leningrad. 

Note. Extremely close to O. amoena C.A.M. and possibly represents 
one of the forms of this highly polymorphic species. 


35. O. hansii Kerner in Nov. pl. sp. decas II (1870) 15 and in Ber. 
naturwiss. Ver. Innsbruck, I (1870) 111.—O. foetida Klotzsch in Reise 
Pr. Wald. Bot. (1862) 111 non Poir. 1786.—O. cernua var. hansii G. 
Beck, Monogr. Orob. (1890) 144, tab. II, f. 33 (4) and in Pflanzenr. 
IV, 261 (1930) 128.—O. inconspicua Gontsch. in Izv. Tadzh. bazy 
AN SSSR, I, 1 (1933) 57.—Ie.: G. Beck, l.c. 

Biennial or perennial (?). Plant up to 45 cm tall, glandular- 
pubescent. Stem yellow, sometimes reddish or violet, middle part up 
to 8 mm thick, slightly thickened at base, with relatively few broadly 
lanceolate, up to 12 mm long, scales. Inflorescence cylindrical, rarely 
lanceolate, usually dense or multiflowered, rarely lax, equaling, shorter 
or somewhat longer than remaining part of stem. Bract scales 
lanceolate, up to 12 mm long. Calyx segments somewhat free, up to 
12 mm long, light brown, often whitish, sometimes with violet tinge, 
ovate at base, lanceolate-subulate, entire or bidentate, with lanceolate- 
subulate teeth. Corolla 15—20 mm long, tubular, distinctly bent forward, 
gradually broadened from constriction toward collar, violet, lighter or 
whitish toward base, sparsely puberulent or glabrous, glabrescent 
outside. Filaments hairy at base, glabrous above, inserted at distance 
of 6-7 mm from base of corolla tube; anthers pubescent along sutures 
of thecae. Style subglabrous. May to July. 

Parasitic usually on species of genus Cousinia Cass.; apparently, 
also on Artemisia L. Rubbly slopes, steppe.—Soviet Central Asia: 
Tien Shan, Pamiro-Alai, mountainous Turkmenia. General distribution: 
Iran, India-Himalayas (northern part), Dzhungaria-Kashgaria (southern 
part), Tibet. Described from India. Type in Innsbriick. 

Note. This species, in form and size of corolla, appears to be 
transitional between O. amoena C.A.M. and O. cernua Loefl.; however, 
in Our opinion it is significantly closer to the former species. Beck 
reported O. hansii Kerner also for Dagestan, probably in error. 


77 


72 


36. O. clarkei Hook. f. Fl. Brit. India, IV (1883) 326; G. Beck, 
Monogr. Orob. (1890) 150 and in Pflanzenr. IV, 261 (1930) 134. 

Biennial or perennial. Plant up to 22 cm tall, glandular-pubescent, 
subglabrous below. Stem yellowish, sometimes with reddish tinge, 
middle part up to 17 mm thick, somewhat thickened at base, with few 
lanceolate, up to 15 mm long, scales. Inflorescence oval, sometimes 
nearly capitate, fairly dense, relatively few-flowered, usually a few 
times as short as remaining part of stem. Bract scales broadly lanceolate, 
up to 18 mm long. Calyx segments free, up to 14 mm long, entire or 
bidentate up to middle, light brown, with faint veins. Corolla 23-28 
mm long, broadly tubular, gradually broadened almost from its base 
towards collar, with constriction, blue in limb, lighter or whitish at 
base, sparsely puberulent outside. Filaments pubescent at base, glabrous 
above, inserted at distance of 3-5 mm from base of corolla tube, 
anthers pubescent along sutures. Style sparsely glandular-pubescent. 
July. 

Parasitic on species of Artemisia L., usually on section Seriphidum 
Bess. Rubbly slopes.— Soviet Central Asia: Pamiro-Alai (Vakhan- 
Inkhashim Region). General distribution: India-Himalayas (northern 
part), Dzhungaria-Kashgaria (southern part), Tibet. Described from 
western Tibet. Type in London. 


37. O. sintenisii G. Beck in Bull. herb. Boiss. 2. sér IV (1904) 
680 and in Pflanzenr. IV, 261 (1930) 134. 

Biennial or perennial (?). Plant 15-20 cm tall, sparsely glandular- 
pubescent, sometimes subglabrous. Stem reddish when dry, slightly 
thickened at base; scales broadly lanceolate, about 10 mm long. 
Inflorescence cylindrical, few flowered, cristate at top, 6-11 cm long. 
Bract scales broadly lanceolate, 1/2 as long as corolla. Calyx segments 
free, entire oblong-lanceolate, long-acuminate, shorter than corolla 
tube. Corolla 30-35 cm long, tubular, considerably though gradually 
broadened from constricted part of tube toward collar, suberect, in 
limb brownish lilac when dry, whitish at base, subglabrous outside. 
Filaments uniformly glandular-pubescent, inserted at distance of 5 mm 
from base of corolla tube. Style sparsely glandular-pubescent. June. 

Host plant not determined. Rubbly slopes.—Caucasus: Perhaps 
found in southern Transcaucasia. General distribution: Armenia- 
Kurdistan. Described from Turkey. Type in Vienna. 

Note. The systematic position of this species, specimens of which 
we did not come across, is not completely certain. Probably, it belongs 
to a separate monotypic series. 


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73 


Series 4. Albescentes Novopokr. in Bot. mat. Gerb. Bot. inst. AN 
SSSR, XII (1950) 284, p.p.—Corolla tube slightly curved, suberect, 
with a slight constriction above the place of insertion of stamens; 
coroila light yellow. Stamens inserted in lower third of corolla tube; 
anthers subglabrous. Plant glandular-pubescent. 


38. O. solenanthi Novopokr. and Pissjauk. in Bot. mat. Gerb. 
Bot. inst. AN SSSR, XII (1950) 284, fig. 6.—Ic.: Novopokr. I.c. 

Perennial or biennial (?). Plant up to 25 cm tall, glandular- 
pubescent. Stem yellowish, often tinged red, middle part about 3 mm 
thick, slightly thickened at base, with few lanceolate, up to 15 mm 
long, scales. Inflorescence short cylindrical or oval, lax, relatively 
few-flowered, usually shorter than remaining part of stem. Bract scales 
broadly lanceolate, up to 15 mm long. Calyx segments free, up to 15 
mm long, whitish, deeply bipartite, teeth lanceolate at base, long- 
subulate. Corolla 20-25 mm long, suberect, broadly tubular, slightly 
constricted above the point of insertion of stamens, light yellow or 
whitish, subglabrous outside. Filaments glabrous, sparsely hairy at 
base, inserted at a height of 6-7 mm from base of corolla tube; anthers 
subglabrous. Style glandular-puberulent only below stigma. June to 
July. 

Parasitic on species of genus Solenanthus Ldb. Forests and scrub.— 
Soviet Central Asia: Pamiro-Alai. Endemic. Described from Hissar 
mountains (Varzob River basin). Type in Leningrad. 


39. O. sulphurea Gontsch. in Izv. Tadzh. bazy AN SSSR, I, 1 
(1933) 53; Novopokr. in Bot. mat. Gerb. Bot. inst. AN SSSR, XII, 
284, fig. 5.—Ic.: Novopokr. l.c. 

Perennial or biennial. Plant up to 65 cm tall, glandular-pubescent. 
Stem yellow, brown when dry, base slightly thickened, set with 
lanceolate, up to 15 mm long, scales; stem usually attached to root of 
host plants in clusters of 2—4. Inflorescence cylindrical or oblong, 
multiflowered, usually lax, equaling or shorter than remaining part of 
stem. Bract scales lanceolate, up to 20 mm long. Calyx segments free, 
10-20 mm long, whitish, bipartite usually up to middle, teeth 
lanceolate-subulate, very rarely entire. Corolla 22-30 mm long, broadly 
tubular, slightly curved forward, suberect, considerably broadened 
toward collar above slight constriction, light yellow, rarely glandular- 
puberulent outside. Filaments hairy at base, glabrous above, inserted 
at height of 6-10 mm from base of corolla tube; anthers sparsely hairy 
along sutures of thecae. Style sparsely puberulent. May to June. 

Host plant, according to N.F. Goncharov, is Crambe kotschyana 
Boiss. Rubbly slopes, scrub.— Soviet Central Asia: Pamiro-Alai. 


Te 


74 


Endemic (?). Described from Tadzhikistan (Vakhsh River valley). Type 
in Leningrad. 

Note. The systematic position of this species is not clear. On the 
basis of floral structure the specimens of this species approach both 
O. amoena C.A.M. and O. kotschyi Reut. and possibly are hybrids. 


Series 5. Cernuae Novopokr. in Bot. mat. Gerb. Bot. inst. AN 
SSSR, XIII (1930) 314.—Corolla tube strongly curved forward, with 
distinct constriction above the place of insertion of stamens, somewhat 
below the middle of corolla tube. Anthers glabrous, rarely subglabrous. 
Parasitic usually on members of the Compositae. 


40. O. cernua Loefl. Iter hisp. (1758) 152; Reut. in DC. Prodr. 
XI, 32; Ldb. Fl. Ross. III, 321; Boiss. Fl. or. 1V, 514.—O. cernua var. 
typica G. Beck, Monogr. Orob. (1890) 14 and in Pflanzenr. IV, 261 
(1930) 122, f. 13 (B); Grossh. Fl. Kavk. IV, 10, p.p.—Ic.: G. Beck, 
l.c. 

Biennial or perennial. Plant 15—30 cm tall, glandular-hairy. Stem 
yellowish or brownish, base slightly thickened, middle part 2-7. mm 
thick, with fairly numerous, broadly lanceolate, 7-12 mm long scales. 
Inflorescence cylindrical or oblong, dense and multiflowered, often 
acuminate at tip, usually more or less equaling remaining part of stem. 
Bract scales with broadly ovate base, lanceolate, 7-13 mm long, often, 
with reddish brown or violet tinge in entire inflorescence. Calyx 
segments free, 7-12 mm long, brownish or reddish brown, usually 
bipartite to less than middle, with lanceolate-subulate teeth, rarely entire. 
Corolla 12-18 mm long, tubular, strongly curved forward and slightly 
patent in limb, whitish toward base, limb bluish or pale violet, 
subglabrous outside. Filaments glabrous or sparsely pubescent at base, 
inserted 5—6 mm above base of corolla tube; anthers glabrous. Style 
subglabrous. May to July. 

Parasitic on members of Compositae (genera Lactuca L., Artemisia 
L., etc.). Rubbly slopes, scrub.—European USSR: (southern part). 
Caucasus: Ciscaucasia, western, eastern and southern Transcaucasia, 
Talysh. Soviet Central Asia: Balkhash Region (southern part), Kara- 
Kum, Syr Darya, Tien Shan, Pamiro-Alai, mountainous Turkmenia. 
General distribution: Mediterranean, Balkan-Asia Minor, Armeina- 
Kurdistan, Iran. Described from Spain. Type in Stockholm. 

Note. The most typical form of this species with completely 


glabrous filaments occurs in southern Crimea and is parasitic on Lactuca 


viminea (L.) Presl. Caucasian and Central Asian plants significantly 
approach O. cumana Wallr. and, apparently, represent several 
physiological races restricted to different genera of the composite 
family. 


80 


i) 


41. O. cumana Wallr. Orob. gen. diask. (1825) 58; Reut. in DC. 
Prodr. XI, 33; Ldb. Fl. Ross. II, 321; Boiss. Fl. or. 1V, 515; Schmalh. 
Fl. II, 298; Gonch. in Sorn. rast. SSSR, IV, 135, f. 421-422; Grossh. 
Opred. rast. Kavk. 354, p.p.—O. arenaria M.B. FI. taur.-cauc. II (1808) 
83 and III (1819) 418, non. Borkh. 1797.—O. bicolor C.A.M. in Ldb. 
Fl. alt. II (1830) 458.—O. cernua var. bicolor Reut. 1.c. 32; Ldb. FI. 
Ross. III, 321.—O. cernua var. cumana G. Beck, Monogr. Orob. (1890) 
143, tab. II, f. 33 (C) and in Pflanzenr. IV, 261 (1930) 128; f. 13 (C); 
Fedch. and Fler. Fl. 891, fig. 860; Grossh. Fl. Kavk. IV, 10; Gonch. 
in Fl. Yugo-Vost. VI, 237, fig..643.—Ic.: Rchb. Ic. Fl. Germ. XX, 
tab. 189; Ldb. Ic. pl. fl. Ross. imp. alt. IV, tab. 390; G. Beck l.c. 
(1890); Fedch. and Fler. 1.c.; Gonch. I.c.; Maltsev. Atlas vazhn. sorn. 
rast. SSSR, I, plate 40.—Exs.: Fl. cauc. exs. No. 350. 

Annual, biennial or perennial. Plant up to 65 cm tall, glandular- 
pubescent. Stem yellowish or brownish, middle part 2-8 mm thick, 
base slightly thickened, with relatively few, broadly lanceolate or ovate 
7-10 mm long scales. Inflorescence cylindrical or oblong, 
multiflowered, usually lax, with distant lower flowers, rarely dense (f. 
densior G. Beck), equaling remaining part of stem or longer. Bract 
scales ovate or ovate-lanceolate acuminate. Calyx segments free, 7—12 
mm long, brownish, often whitish in lower part, usually bipartite, less 
than middle, with lanceolate-subulate teeth or entire. Corolla 12—20 
mm long, tubular, strongly curved forward and slightly patent in limb, 
base whitish, limb bluish or whitish, violet, usually paler colored than 
in O. cernua Loefl., subglabrous outside. Filaments subglabrous, 
sparsely pubescent at base, inserted 5-7 cm above base of corolla 
tube; anthers glabrous, rarely sparsely hairy. Style subglabrous. May 
to August (Plate V, fig. 3). 

Parasitic on composites (predominantly on species of genera 
Artemisia L., Helianthus L., Xanthium L., Karelinia Less., etc.), less 
commonly on Solanaceae (Lycopersicum Mill. and Nicotiana L.). 
Steppe, rubbly slopes, often in fields and gardens, roadsides, weedy 
habitats. — European USSR: Middle Dnieper, Volga-Don, Trans-Volga 
Bessarabia, Black Sea Region, Lower Volga, Crimea. Caucasus: All 
regions. Western Siberia: Upper Tobol, Irtysh, Altai. Eastern Siberia: 
Dauria (southern part). Soviet Central Asia: All regions. General 
distribution: Central Europe (southern part), Mediterranean, Balkans- 
Asia Minor, Armenia-Kurdistan, Iran, Dzhungaria-Kashgaria, Mongolia. 
Described from Astrakhan Region. Type was in Berlin. 

Note. A polymorphic species, differentiating into a large number 
of morphologically slightly distinct, as yet insufficiently studied, 
physiological races, partly transitional to the foregoing species. 


81 


76 


Economic importance. It is a serious pest of sunflower, sometimes 
growing in abundance. It is possible to distinguish physiological races 
of this species, arbitrarily denoted as A and B, of which the latter, 
evidently, is relatively recently evolved and so-called ‘vicious’ 
Orobanche, which is able to parasitize many varieties of sunflower; 
race A is not as harmful. Much less frequently it parasitizes other 
cultivated plants: tomato, tobacco, safflower, perilla, zinnia. 


42. O. brevidens Novopokr. in Bot. mat. Gerb. Bot. inst. AN 
SSSR, XII (1950) 282, fig. 4.—Ic.: Novopokr. l.c. 

Biennial or perennial. Plant 20-45 cm tall, glandular-pubescent. 
Stem yellowish or brownish, often tinged red, middle part up to 12 
mm thick, slightly thickened at base, with fairly numerous, broadly 
lanceolate, up to 20 mm long scales. Inflorescence cylindrical, 
multiflowered, fairly dense, more or less equaling remaining part of 
stem. Bract scales ovate or ovate-lanceolate, up to 15 mm long. Calyx 
segments from broadly ovate at base to lanceolate, entire, rarely shortly 
bidentate at tip, up to 15 mm long, light brown, often whitish with 
faint veins. Corolla 20-25 mm long, distinctly curved forward and 
slightly patent in limb, tubular, yellowish, whitish toward limb, 
subglabrous outside. Filaments sparsely hairy at base, subglabrous, 
inserted 6-7 mm above base of corolla tube; anthers glabrous. Style 
glabrous. June to July. 

Host plant not determined; reported on species of Alhagi, but this 
information needs confirmation. 

Steppe, alkali soils —Soviet Central Asia: Aralo-Caspian 
(southeastern part), Kyzyl Kum. Endemic. Described from Kazakhstan 
(between Kzyl-Orda and landmark Bish-Arak). Type in Leningrad. 

Note. This species is one of the physiological races of series 
Cernuae and is very close to certain forms of O. cumana Wallr. In our 
opinion, its resemblance to O. amoena C.A.M., as believed by I.V. 
Novopokrovsky, is without basis. 


43. O. solmsii Clarke in Hook. f. Fl. Brit. Ind. IV (1883) 325; G. 
Beck, Monogr. Orob. (1890) 147 and in Pflanzenr. IV, 261 (1930) 
129.—(. nikitinii Novopokr. in Bot. mat. Gerb. Bot. inst. AN SSSR, 
XII (1950) 287, fig. 7.—O. mulgedii Novopokr. |.c. XIII (1950) 314.— 
Ic.: Novopokr. l.c. 

Biennial or perennial. Plant up to 35 cm tall, glandular-pubescent. 
Stem yellowish, middle part 3-5 cm thick, thickened at base; scales 
lanceolate or ovate-lanceolate, up to 15 mm long. Inflorescence 
cylindrical or oblong, multiflowered, and fairly dense, equaling 
remaining part of stem or somewhat shorter. Bract scales ovate or 


8 


N 


aT 


ovate-lanceolate, almost equaling calyx. Calyx segments free, 7—12 
mm long, entire or bipartite up to middle, with lanceolate-subulate 
teeth, light brown or whitish, with faint veins. Corolla (12)14—18(20) 
mm long, tubular, distinctly bent forward and slightly patent in limb, 
yellowish, often whitish at base, subglabrous outside. Filaments 
glabrous or sparsely pubescent at base, inserted 5-7 mm above base 
of corolla tube; anthers glabrous or subglabrous. Style glabrous or 
subglabrous. May-July. 

Parasitic on composites; reported on Lactuca tatarica L. and 
species of Artemisia L. Rubbly slopes, sandy soils.—Soviet Central 
Asia: mountainous Turkmenia, Pamiro-Alai. General distribution: Iran, 
India-Himalayas (northern part), Dzhungaria-Kashgaria (southern part). 
Described from western Himalayas. Type in London. 

Note. There are hardly any distinctive morphological differences 
between the foregoing species O. solmsii Clarke and its synonyms, 
although O. nikitinii is close to O. amoena C.A.M., according to N. 
Novopokrovsky (in our opinion, without sufficient basis). All 
considered, O. solmsii probably represents a color variety of O. cernua 
Loefl. 


44. O. badchysensis Novopokr. and V. Nikitin in Bot. mat. Gerb. 
Bot. inst. AN SSSR, XIII (1950) 315. 

Biennial or perennial (?). Plant up to 15 cm tall, glandular- 
pubescent. Stem, about 4 cm long excluding inflorescence, base slightly 
thickened; lower scales shorter, ovate, upper ones longer, ovate- 
lanceolate. Inflorescence cylindrical, almost equaling remaining part 
of stem, 2.2-2.5 mm broad, rounded at tip. Bract scales ovate, obtuse, 
1/2 as long as flowers, glandular-pubescent along midrib and margin. 
Calyx segments free 1/2—2/5 as long as corolla, fairly broad, glandular- 
puberulent, parted up to middle or somewhat deeper into 2 deltoid- 
lanceolate teeth, each with 3 veins. Corolla 12-15 mm long, tubular, 
fairly strongly curved forward, subglabrous (less noticeable, glandular- 
puberulent, in upper part), yellowish-dun colored when dry, whitish in 
middle and lower parts. Filaments in lower part sparsely hairy, inserted 
somewhat above middle of corolla tube; anthers glabrous. Style 
glabrous. May to June. 

Parasitic on Chesneya kopetdaghensis Boriss. Rubbly slopes.— 
Soviet Central Asia: mountainous Turkmenia (Badkhyz). General 
distribution: Probably Iran. Described from Kopet-Dag. Type in 
Leningrad (?), isotype in Ashkhabad. 

Note. We have not seen specimens of this species, and its systematic 
position is not fully understood. Apparently, it is so close to O. solmsii 
Clarke that the original diagnosis does not reveal any essential 
differences. 


8 


WwW 


78 


Series 6. Camptolepides Tzvel.—Corolla tube suberect, with less 
noticeable constriction. Stamens inserted somewhat below middle of 
corolla tube; anthers subglabrous. Bract scales broadly ovate to 
suborbicular, tip usually attenuated, with short, glabrous curved cusp. 
Parasitic on buckwheat family Polygonaceae Linrl. 


45. O. camptolepis Boiss. and Reut. in Boiss. Fl. or. IV, 515; G. 
Beck, Monogr. Orob. 145.—O. cernua var. latebracteata G. Beck in 
Pflanzenr. IV, 261 (1930) 126.—O. gontscharovii Novopokr. in Bot. 
mat. Gerb. Bot. inst. AN SSSR, XIII (1950) 316.—O. schugnanica 
Novopokr. I.c. 319.—Ic.: Novopokr. I.c. 320. 

Biennial or perennial. Plant 8-20 cm tall, glandular-pubescent, 
sometimes subglabrous. Stem yellowish, often tinged red, middle part 
up to 10 mm thick, base somewhat thickened; scales fairly numerous 
up to 10 mm long, broadly ovate to suborbicular, thick, brownish; tip 
often attenuated with short, lanceolate cusp, hooked when dry. 
Inflorescence oblong or short cylindrical, fairly dense, equaling 
remaining part of stem or somewhat longer. Bract scales up to 10 mm 
long, resembling scales of stem, usually accumbent toward flowers, or 
midrib densely gray-puberulent. Calyx segments free, up to 10 mm 
long, fairly thick, light brown, bidentate up to half its length, with 
lanceolate-subulate teeth. Corolla 12-18 mm long, slightly declinate 
from inflorescence axis, tubular, suberect, slightly patent in limb, pale 
violet or light blue (?), base whitish, usually pinkish when dry, rarely 
glandular-pubescent outside. Filaments glabrous or with few hairs at 
base, inserted 4-6 mm above base of corolla tube; anthers and style 
subglabrous. June to August (Plate V, fig. 4). 

Parasitic on small shrubs and subshrubs of family Polygonaceae; 
reported on species of Atraphaxis L., Polygonum paronychioides 
C.A.M. and P. maritimum L. Rubbly slopes, steppe.— Soviet Central 
Asia: Tien Shan, Syr Darya, Pamiro-Alai. General distribution: Near 
East, Iran, Dzhungaria-Kashgaria. Described from Turkey. Type in 
Geneva. 

Note. A form (O. cernua var. laterbracteata f. nanantha G. Beck) 
close to this species has been reported by Beck from Crimea, possibly 
mistakenly. 


Series 7. Crassicaules Novopokr. in Boi. mat. Gerb. Bot. inst. AN 
SSSR, XIII (1950) 317.—Corolla tube slightly bent, with or without 
a distinct constriction above the point of insertion of stamens. Stamens 
inserted somewhat below middle of corolla tube, rarely in its lower 
third; anthers hairy along sutures of thecae, rarely glabrous. Stem usually 
clavate at base. Parasitic on carrot family. 


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79 


46. O. kotschyi Reut. in DC. Prodr. XI (1847) 33; Boiss. FI. or. 
IV, 514; G. Beck, Monogr. Orob. 146, p.p. tab. II, f. 35 (1) and in 
Pflanzenr. IV, 261, 129, p.p. f. 13 (D).—O. spectabilis Reut. apud 
Boiss. et Reut. in Nouv. Mém. Soc. Nat. Mosc. XII (1860) 169; Boiss. 
l.c. 514.—O. bucharica Novopokr. in Bot. mat. Gerb. Bot. inst. AN 
SSSR, XIII (1950) 318.—O. xanthostachya Novopokr. |.c. 321.—Ic.: 
G. Beck, l.c. 

Biennial or perennial. Plant up to 60 cm tall, glandular-pubescent, 
usually with mixture of longer whitish hairs or wholly white arachnoid- 
lanate in inflorescence. Stem yellowish, middle part 5—15 mm thick, 
clavate at base (up to 4 cm thick), covered with lanceolate or ovate- 
lanceolate, up to 10 mm long, scales. Inflorescence cylindrical or 
oblong, fairly dense and multiflowered, more or less equaling remaining 
part of stem. Bract scales ovate-lanceolate or broadly lanceolate, 
somewhat longer than or equaling calyx, usually more or less whitish- 
lanate on midrib. Calyx segments free, 12-15 mm long, light brown, 
with faint, darker veins, usually incised to more than middle into 2 
lanceolate-subulate teeth, rarely short bidentate. Corolla 18-25 mm 
long, tubular, slightly bent forward, with distinct constriction above 
place of insertion of stamens, slightly patent in limb, brownish yellow, 
whitish toward base, rarely with extremely pale blue tinge in limb, 
which rapidly disappears on drying, glandular-hairy outside. Filaments 
sparsely hairy at base, glabrous above, inserted 7-10 mm above base 
of corolla tube; anthers pubescent at base. Style subglabrous. May to 
August. 

Parasitic on umbellifers; observed on species of genera Ferula L., 
Prangos L., Zozimia Hoffm., etc. Rubbly slopes, scrub.— Soviet Central 
Asia: mountainous Turkmenia, Pamiro-Alai. General distribution: 
Armenia-Kurdistan, Iran, Dzhungaria-Kashgaria. Described from 
southern Iran. Type in Paris. 

Note. A polymorphic species, highly variable in the degree of 
pubescence, types of inflorescence, size of corolla and other features. 
However, clear adaptation of individual forms to different host plants 
does not seem to have been observed. It is possible to recognize a 
whitish, arachnoid-lanate form of this species—f. spectabilis (Reuter) 
G. Beck [in Pflanzenr. IV, 261 (1930) 130], characterized, besides, by 
an especially long and multiflowered inflorescence with darker colored 
flowers in general. Specimens with smaller flowers and weak 
pubescence are not always clearly distinguishable from O. solmsii 
Clarke. 


47. O. gigantea (G. Beck) Gontsch. in Izv. Tadzh. bazy. AN SSSR, 
I, 1 (1933) 55.—O. kotschyi var. gigantea G. Beck, Monogr. Orob. 
(1890) 147, tab. II, f. 35 (2) and in Pflanzenr. IV, 261 (1930) 131.— 


85 


80 


O. pamiroalaica Gontsch. and Novopokr. in Bot. mat. Gerb. Bot. inst. 
AN SSSR, XII (1950) 277.—Ic.: G. Beck, l.c. (1890). 

Biennial or perennial. Plant up to 65 mm tall, glandular-pubescent, 
usually with mixture of longer, whitish hairs in inflorescence. Stem 
yellowish, middle part 5-20 mm thick, clavate at base (up to 5 cm 
thick), covered with lanceolate, up to 25 mm long, scales. Inflorescence 
cylindrical, oblong or oval, fairly dense, usually more or less equaling 
remaining part of stem. Bract scales lanceolate, up to 22 mm long, 
white pubescent or lanate on midrib. Calyx segments free, 12—20 cm 
long, light brown, usually incised more than middle into 2 lanceolate- 
subulate teeth, rarely short bidentate or even entire. Corolla 25—40 cm 
long, tubular, slightly bent forward to almost straight, with less 
noticeable constriction, often distinctly patent towards limb, brown- 
yellow to whitish at base, glandular hairy outside. Filaments hairy at 
base, glabrous above, but often glandular-pubescent below anthers, 
inserted 10-14 mm above base of corolla tube; anthers hairy at base. 
Style rarely with glandular hairs. May to August. 

Parasitic on umbellifers; observed on species of Ferula L. and 
Libanotis Cr. Rubbly slopes, scrub.— Soviet Central Asia: Tien Shan, 
Syr Darya, Pamiro-Alai. General distribution: Dzhungaria-Kashgaria. 
Described from Tadzhikistan. Type in Leningrad. 

Note. Of the majority of forms of this polymorphic species (which 
is very close to the foregoing species), we should mention f. talassica 
Tzvel. f. nova, from western Tien Shan, having an oval, sometimes 
almost capitate and few-flowered inflorescence and almost straight 
flowers. Its corolla resembles O. amoena C.A.M. in shape, just like 
many other large-flowered plants of O. gigantea. It is not surprising 
that several specimens of this last species were identified by Beck as 
“O. amoena f. colossea G. Beck.” Apparently, O. pamiroalaica, was 
described from an atypical specimen of O. gigantea. 


48. O. sordida C.A.M. in Ldb. FI. alt. IT (1830) 455; Reut. in DC. 
Prodr. XI, 31; Ldb. Fl. Ross. HI, 320; G. Beck, Monogr. Orob. 167, 
tab. III, f. 44 and in Pflanzenr. IV, 261, 242.—O. irtyschensis 
Sergievsk. in Sist. zam. po mat. gerb. Tomsk. Univ. Nos. 1—2 (1939) 
4 and in Kryl. Fl. Zap. Sib. X, 2539.—Ic.: Ldb. Ic. pl. fl. ross. imp. 
alt. IV, tab. 386; G. Beck, l.c. 

Biennial or perennial. Plant up to 50 cm tall, white hairy. Stem 
brownish yellow, sometimes tinged red, middle part up to 10 mm 
broad, base usually clavate, covered with lanceolate scales up to 15 
mm long. Inflorescence cylindrical, fairly dense, multiflowered, 
equaling somewhat longer than remaining part of stem. Bract scales 
lanceolate. Calyx segments free, up to 12 mm long, light brown, 


86 


81 


bidentate to less than middle or entire, lanceolate-subulate at tip. 
Corolla 16-25 mm long, tubular, almost straight with indistinct 
contriction above point of insertion of stamens, slightly expanded 
toward limb, brownish yellow, whitish toward base, glandular- 
pubescent outside. Filaments hairy at base, glabrous or subglabrous 
above, inserted 7-9 mm above base of corolla tube; anthers hairy 
along sutures of thecae. Style rarely glandular-puberulent. June to 
July (Plate V, fig. 5). 

Host plant not determined probably parasitizes umbellifers. Rubbly 
slopes, scrub.— Western Siberia: Altai (southern part). Soviet Central 
Asia: Balkhash Region (eastern part), Dzhungaria-Tarbagatai, Tien 
Shan. General distribution: Dzhungaria-Kashgaria. Described from 
eastern Kazakhstan (vicinity of Lake Zaisan). Type in Leningrad. 


49. O. vitellina Novopokr. in Grossh. Opred. rast. Kavk. (1949) 
354 and in Bot. mat. Gerb. Bot. inst. AN SSSR, XII (1950) 288, fig. 
8.—Ic.: Novopokr. l.c. 

Biennia! or perennial. Plant up to 30 cm tall, white hairy. Stem 
brownish yellow, up to 5 mm thick in middle part, clavate at base, 
with fairly numerous narrowly lanceolate, up to 15 mm long, scales. 
Inflorescence cylindrical, fairly dense, many-flowered, equaling or 
somewhat shorter than remaining part of stem. Bract scales lanceolate, 
up to 12 mm long, midrib densely white hairy. Calyx segments free, 
up to 12 mm long, usually parted up to middle into 2 lanceolate- 
subulate teeth; light brown, with very weak veins. Corolla 18-22 mm 
long, tubular, almost straight, slightly broadened toward limb, 
yellowish, whitish toward base, rarely glandular-puberulent outside. 
Filaments subglabrous, sparsely pubescent at base, inserted 5-7 mm 
above base of corolla tube; anthers densely hairy along sutures of 
thecae. Style subglabrous. May to June. 

Host plant not determined; probably parasitizes umbellifers. Rubbly 
slopes, scrub.—Caucasus: Southern Transcaucasia. General 
distribution: probably Armenia-Kurdistan. Described from Nachichevan 
ASSR. Type in Leningrad. 


50. O. glabricaulis Tzvel. sp. n. in Addenda XXII, 685. 

Biennial or perennial. Plant up to 35 cm tall, glabrous. Stem 
brownish-yellow, up to 15 mm thick in middle part, clavate at base 
(up to 3 cm thick), with relatively few broadly lanceolate, up to 15 
mm long, scales. Inflorescence short cylindrical, tairly dense, many- 
flowered, usually slightly shorter than remaining part of stem. Bract 
scales lanceolate, up to 18 mm long. Calyx segments free, 8-12 mm 
long, ovate at base, lanceolate, entire or incised less than middle into 


87 


82 


2 lanceolae teeth, light brown, with weak veins. Corolla 18-25 mm 
long, almost straight, slightly inflated below place of insertion of 
stamens, slightly spreading out in limb, yellowish brown, whitish at 
base, glabrous. Filaments glabrous, inserted 5-7 mm above base of 
corolla tube. Anthers and style glabrous. May to June. 

Clayey, saline slopes.— Caucasus: Southern Transcaucasia (in 
vicinity of the city of Nachichevan). General distribution: Probably 
Armenia-Kurdistan. Described from Nakhichevan ASSR (Nachitchevan, 
in argillosis salsis, 27 V 1934, L. Prilipko). Type in Leningrad, cotype 
in Baku. 

Note. Differs from other species of the series in its total absence 
of pubescence; and from O. vitellina Novopokr., in having a 
considerably thicker stem. Probably belongs to a separate series. 


Subsection 2. Angustatae G. Beck in Hal. u. Braun, Nachr. 
Zur Fl. Nieder-Oesterr. 125; Monogr. Orob. 152 and in Pflanzenr. IV, 
261, 136.—Corolla with relatively short tube, often campanulate, 
spreading out from the place of insertion of stamens toward throat, 
narrowed toward base. Stamens inserted close to base of corolla tube, 
rarely in its lower third, extremely rarely close to its middle (several 
species of group Minores G. Beck). 

Note. We accept the proposal of Beck regarding the fairly natural 
division of this subsection into groups (grex), approximately 
corresponding to series. 


Series 1. Galeatae Beck in Hal. u. Braun, Nauchr. zur Fl. Nieder- 
Oesterr. (1882) 125; Monogr. Orob. 134 and in Pflanzenr. IV, 261, 
222.—Corolla 18-30 mm long, usually obliquely erect, with relatively 
long tube, barely sinuate, galeate upper lip, its oblong dorsal line 
straight to a considerable extent. Calyx segments broad, bidentate, 
non-subulate at tip. Pubescence brownish or yellowish. 


51. O. gamosepala Reut. in DC. Prodr. XI (1847) 15; Ldb. FI. 
Ross. III, 315; Boiss. Fl. or. IV, 503; G. Beck, Monogr. Orob. 154, 
tab. Il, f. 41 and in Pflanzenr. IV, 261, 225, f. 16 (E); Grossh. FI. 
Kavk. IV, 11.—Ic.: G. Beck, l.c. 

Perennial. Plant 5-45 cm tall, sparsely glandular-pubescent, 
sometimes subglabrous. Stem reddish-brown, somewhat thickened at 
base, with few oblong-ovate, obtuse, up to 20 mm long, scales. 
Inflorescence cylindrical or oval, lax or dense in upper part, often 
relatively few-flowered, usually shorter than remaining part of stem. 
Bract scales ovate, acuminate, nearly equaling calyx. Calyx 
gamosepalous, rarely incised up to base towards inflorescence axis, 


88 


83 


with 2-4 lanceolate teeth. Corolla 25-35 mm long, obliquely erect at 
tip, with relatively long tube, gradually broadening toward limb, dark 
red, rarely brownish red, rarely glandular-puberulent outside, its 
longitudinal dorsal line erect for a considerable extent, upper lip barely 
sinuate, carinate, galeate. Filaments subglabrous, inserted 3—7 mm 
above base of corolla tube; anthers more or less hairy. Style subglabrous. 
June to August (Plate V, fig. 6). 

Parasitic on species of Geranium L. Subalpine meadows, rubbly 
slopes of upper mountainous belt.— Soviet Central Asia: All regions 
besides Talysh. General distribution: Probably Armenia-Kurdistan. 
Described from Caucasus (Mt. Mashuk). Type in Geneva, isotype in 
Leningrad. 


52. O. lutea Baumg. Enum. stirp. Transsylv. II (1816) 215 excl. 
syn.; G. Beck, Monogr. Orob. 163, tab. III, f. 44 and in Pflanzenr. IV, 
261, 237, f. 16 (J—N); Fedch. and Fler. Fl. Evrop. Ross. 892, fig. 863; 
Grossh. Fl. Kavk. IV, 11.—0O. rubens Wallr. Schrd. crit. I (1822) 307 
and Orob. gen. diask. (1825) 46; Reut. in DC. Prodr. XI, 25; Ldb. FI. 
Ross. III, 318; Boiss. Fl. or. IV, 511; Schmalh. Fl. II, 297.—Ic.: 
Rchb. Ic. Fl. Germ. XX, tab. 171; G. Beck, |.c.; Fedch. and Fler. l.c.; 
Hegi, Ill. Fl. VI, f. 83 and 84 (a—d). 

Perennial. Plant 20—60 cm tall, densely glandular-pubescent. Stem 
yellow or brownish, middle part 3-8 mm thick, somewhat thickened 
at base, with fairly numerous lanceolate, 12-25 mm long, scales. 
Inflorescence cylindrical, rarely oval, fairly dense and many-flowered, 
sometimes lax, usually shorter than remaining part of stem. Bract 
scales broadly lanceolate, 12-18 mm long. Calyx segments free, 7—12 
mm long, brownish and fairly thick with distinct veins, extremely 
broad at base, split up to middle or more into 2 broad patent lanceolate 
unequal teeth, anterior tooth normally shorter. Corolla 18—22 mm long, 
obliquely erect, rarely almost horizontally inclined to axis, with 
relatively long tube, gradually spreading out at limb, brownish or 
yellow, limb sometimes with pinkish tinge, densely glandular-pubescent 
outside, its longitudinal dorsal line straight to a considerable extent, 
upper lip entirely or barely sinuate, galeate. Filaments hairy along one- 
third their length from base, glabrous above, inserted 4-5 mm above 
base of corolla tube; anthers pubescent at base. Style subglabrous. 
May to July (Plate V, fig. 7). 

Parasitic predominantly on perennial species of Medicago L., rarely 
on other legumes. Rubbly slopes, steppe, scrub, occasionally in fields. — 
European USSR: Upper Dnieper, Middle Dnieper, Bessarabia, Black 
Sea Region, Lower Don, Crimea. Caucasus: All regions. Soviet Central 
Asia: mountainous Turkmenia, Pamiro-Alai. General distribution: 


89 


84 


Central and Atlantic Europe, western Mediterranean, Balkans-Asia 
Minor, Armenia-Kurdistan, Iran. Described from Romania. Type in 
Vienna. 

Note. One specimen from Dagestan has the corolla strongly bent 
forward at the point of insertion of the stamens and almost horizontally 
divergent from the axis of stem. This form is referred to as var. 
buekiana Koch [Syn. fl. Germ. ed. 2 (1844) 619]. Such plants 
occasionally occur throughout the entire range of O. lutea and, 
evidently, represent atypical or hybrid forms. 

Economic importance. Sometimes harmful to alfalfa and clover, 
when it grows abundantly in the fields of such crops. For control, see 
Economic Importance of the genus. 


53. O. linezevskyi Novopokr. in Bot. mat. Gerb. Bot. inst. AN 
SSSR, XIII (1950) 327. 

Perennial. Plant up to 60 cm tall, densely glandular-pubescent. 
Stem yellowish brown, often tinged red, middle part up to 10 mm 
thick, base clavate (up to 4 cm thick) and densely covered with 
lanceolate, thick scales, above with more rare, lanceolate or ovate- 
lanceolate, up to 20 mm long, scales. Inflorescence cylindrical, fairly 
thick in upper part, lax below, with distant flowers, equaling remaining 
part of stem or shorter. Bract scales broadly lanceolate, up to 20 mm 
long. Calyx segments free, up to 15 mm long, brownish and fairly 
thick, bidentate from broadly ovate base, teeth narrowly lanceolate, 
gradually acuminate, usually almost equaling connate part of segments. 
Corolla 20-25 mm long, with relatively long tube, somewhat inflated 
at place of insertion of stamens and sharply bending forward, almost 
horizontally inclined to axis of inflorescence, yellowish brown when 
dry, glandular-pubescent outside, its longitudinal dorsal line almost 
straight in middle; upper lip barely sinuate, galeate. Filaments hairy at 
base, glabrous above, but sparsely glandular-pubescent below anthers, 
inserted 4-5 mm above base of corolla tube; anthers and style 
subglabrous. Stigma usually yellow. June to July. 

Parasitic on legumes; observed on Medicago sativa L. Rubbly 
slopes, scrub.— Soviet Central Asia: Pamiro-Alai. Endemic (7). 
Described from southern Tadzhikistan. Type in Leningrad. 

Note. A. doubtful species, probably described from a large 
specimen of O. lutea var. buekiana Koch. mentioned above. This is 
confirmed by collections of normal specimens of O. lutea Baumg. 
from the same locality. 


54. O. teucrii Holand. Exs. (1824) and FI. de la Moselle éd. 1, II 
(1829) 322; Reut. in DC. Prodr. XI, 21; ? Boiss. Fl. or. IV, 508; 


90 


85 


G. Beck, Monogr. Orob. 161, tab. III; f. 43 and in Pflanzenr. IV, 261, 
234.—Ic.: Rchb. Ic. Fl. Germ. XX, tab. 69; G. Beck, |.c.; Hegi, Ill. Fl. 
VI, f. 82 (g-i). 

Perennial. Plant up to 25 cm tall, glandular-pubescent (brownish 
pubescence). Stem brownish yellow, often reddish colored; middle 
part up to 4-5 mm thick, slightly thickened at base, with fairly 
numerous lanceolate, up to 15 mm long, scales. Inflorescence oblong, 
lax and often few-flowered, usually shorter than remaining part of 
stem. Bract scales broadly lanceolate, up to 15-20 mm long. Calyx 
segments free, rarely connate in front, up to 10-12 mm long, brownish, 
bidentate from broadly ovate base to half its length or less, with 
narrowly lanceolate teeth. Corolla 18-22 mm long, inclined obliquely 
upward, with relatively long tube, gradually broadening toward limb, 
brownish lilac, often tinged red, usually dark brown when dry, 
glandular-pubescent outside, its longitudinal dorsal line straight to a 
considerable extent. Filaments hairy up to 1/3 length, glabrous above 
but glandular-puberulent below anthers, inserted 4-5 mm above base 
of corolla tube; anthers subglabrous. Style glandular-pubescent. Stigma 
red. June to July. 

Parasitic on species of Teucrium L. Rubbly slopes.—European 
USSR: Upper Dnieper (Carpathian mountains). General distribution: 
Central and Atlantic Europe, western Mediterranean, Balkans. 
Described from France. Type in Paris. 


55. O. schelkovnikovii Tzvel. sp. n. in Addenda XXI, 685. 

Perennial or biennial. Plant up to 45 cm tall, diffusely glandular- 
pubescent. Stem brownish or reddish yellow; middle part up to 1 cm 
thick, base somewhat thickened, lower part subglabrous, with relatively 
few broadly lanceolate, up to 20 mm long, scales. Inflorescence 
cylindrical or oblong, many-flowered, moderately dense, nearly 
equaling remaining part of stem. Bract scales broadly lanceolate, up 
to 18 mm long, densely glandular-pubescent on back of midrib. Calyx 
segments (on side of bract scales) largely connate in front, 10-15 mm 
long, diffusely glandular-pubescent, brownish yellow, incised almost 
up to middle into 2 lanceolate teeth. Corolla 24-30 mm long, tubular- 
campanulate, considerably broadened at throat, brownish lilac when 
dry, diffusely glandular-pubescent outside, glabrous or subglabrous 
inside; longitudinal dorsal line in middle part of corolla tube almost 
straight, arcuate toward base and limb; upper lip sinuate, with very 
broad reflexed lobes; lower lip with 3 suborbicular lobes; all lobes 
irregularly, serrulate along margin, diffusely glandular-pubescent. 
Filaments hairy near base above, glandular-puberulent below anthers, 
inserted 6—9 mm above base of corolla tube; anthers pubescent along 


9 


— 


86 


sutures of thecae. Style glandular-puberulent in upper part. Stigma 
yellow. July to August. 

Host plant not determined. Subalpine meadows.— Caucasus: 
Southern Transcaucasia. General distribution: Probably, Armenia- 
Kurdistan. Described from Armenia (Mt. Agmagan). Type in Yerevan. 

Note. The species is described from the collection of A. 
Shelkovnikov from subalpine meadows of the Agmagan Mountains in 
Armenia (Flora Armeniae, No. 1893, mont. Agmagan, prat. subalpin. 
Jelidja, 23-25 VII 1926, A. Shelkovnikov). It occupies a fairly 
independent position in the group Galeatae, differing from O. vulgaris 
Poir. in the extremely high insertion of the stamens in the corolla tube; 
from O. lutea Baumg. in the color and shape of the corolla and connate 
calyx segments; from O. teucrii Holand. in the distinct veins on the 
calyx segments and broader corolla lobes. 


56. O. vulgaris Poir. in Lam. Encycl. IV (1797) 621; Gonch. in 
Fl. Yugo-Vost. VI, 619.—O. caryophyllacea Smith in Trans. Linn. 
Soc. IV (1797) 169; M.B. FI. taur.-cauc. II, 81; G. Beck, Monogr. 
Orob. 155, tab. II, f. 42 and in Pflanzenr. IV, 261; 226, f. 16 (F—H); 
Schmalh. Fl. II, 295; Fedch. and Fler. Fl. 891, fig. 861 and 862; 
Grossh. Fl. Kavk. IV, 11.—O. galii Duby. Bot. Gall. I (1828) 349; 
Reut. in DC. Prodr. XI, 20; Ldb. Fl. Ross. III, 317; Boiss. Fl. or. IV, 
508.—? O. quadrifida C. Koch in Linnaea, XXII (1849) 665; Boiss. 
l.c. 503; Grossh. Fl. Kavk. IV, 11.—O. buhsei Reut. apud. Boiss. and 
Buhse. in Mém. Soc. Nat. Mosc. XVIII (1860) 169.—Ic.: G. Beck, 
l.c.; Fedch. and Fler. I.c.; Hegi, Ill. Fl. VI, f. 82 (d-f). 

Perennial. Plant 15-40 cm tall, usually brownish, fairly densely 
glandular-pubescent. Stem yellowish brown, often tinged red or violet, 
middle part up to 7 mm thick, base slightly thicker, covered with 
fairly numerous, lanceolate, up to 25-30 mm long, scales. Inflorescence 
short-cylindrical or oval, often lax and few-flowered, usually 
considerably shorter than remaining part of stem. Bract scales ovate- 
lanceolate, up to 30 cm long. Calyx segments free or connate in front, 
up to 15 (rarely 22) mm long, light brown, at base extremely broad 
membranous, usually incised up to middle into 2 unequal lanceolate 
teeth, rarely subentire. Corolla 20-30 mm long, inclined obliquely 
upward, with relatively long, though broad tube, almost campanulate, 
brownish lilac, usually tinged red, densely glandular-pubescent outside, 
its longitudinal dorsal line straight to considerable extent; upper lip 
barely sinuate, galeate. Filaments hairy for 1/4—1/3 of length from 
base, glabrous above but glandular-puberulent below anthers, inserted 
at 1.5-3 mm, rarely up to 5 mm above base of corolla tube; anthers 


92 


87 


glabrous or subglabrous. Style sparsely glandular-hairy; stigma usually 
dark red. May to July. 

Parasitic on madder family (Rubiaceae Juss.), usually on species 
of Galium L. Rubbly slopes, steppe, scrub.— European USSR: Upper 
Dniester, Upper Dnieper, Middle Dnieper, Volga-Don, Bessarabia, 
Black Sea Region, Lower Don, Crimea. Caucasus: All regions. Soviet 
Central Asia: Kara-Kum, mountainous Turkmenia. General 
distribution: Scandinavia, Central and Atlantic Europe, western 
Mediterranean, Balkans-Asia Minor, Armenia-Kurdistan, Iran. 
Described from France. Type in Paris. 

Note. In the Nakhitchevan SSSR, plants with a higher (up to 5 mm 
from base of corolla tube) insertion of the stamens have been 
observed.—var. buhsei (Reuter) G. Beck [Monogr. Orob. (1890) 160]. 
Probably, they represent an independent physiological race. O. 
quadrifida C. Koch, described from Georgia, apparently, from 
specimens of O. vulgaris Poir. with the calyx segments connate in 
front. 


Series 2. Glandulosae G. Beck in Hal. u. Braun. Nachtr. zur FI. 
Nieder-Oesterr. (1882) 132; Monogr. Orob. 135 and in Pflanzenr. IV, 
261, 142.—Corolla with short and broad tube, usually campanulate, 
15-25 mm long, usually glandular-pubescent outside, with mixture of 
dark colored (red or violet) glandular hairs, which can be recognized 
in dry material from their black nodules (points) at their bases, its 
longitudinal dorsal line almost straight in middle part, rarely almost 
uniformly curved. Calyx segments entire, rarely bidentate. Pubescence 
short, brownish or yellowish. 


57. O. hymenocalyx Reut. in DC. Prodr. XI (1847) 20; Ldb. FI. 
Ross. III, 317; G. Beck, Monogr. Orob. 214 and in Pflanzenr. IV, 261, 
167; Grossh. Fl. Kavk. IV, 12. 

Perennial. Plant up to 40 cm tall, glandular-pubescent. Stem 
brownish when dry, with relatively few oblong-lanceolate, up to 22 
mm long, scales. Inflorescence cylindrical, sufficiently dense, up to 
17 cm long. Bract scales from oval at base to narrowly lanceolate, 
exceeding flower, densely glandular-hairy. Calyx segments connate in 
front, extremely broad at base, bidentate; teeth short, subdeltoid, shorter 
than connate part of segments. Corolla broadly campanulate, about 20 
mm long, brownish when dry, glandular-pubescent outside, its 
longitudinal dorsal line almost straight in middle part; upper lip more 
or less bifid, barely galeate, with broad, orbicular lobes. Filaments 
glabrous or subglabrous, inserted 2-3 mm above base of corolla tube; 


93 


88 


anthers sparsely pubescent. Style glandular-pubescent; stigma yellow. 
June to July (?). 

Host plant not determined. Rubbly slopes.— Caucasus: Eastern 
Transcaucasia. Endemic. Described from Kirovabad Region. Type in 
Geneva. 

Note. A doubtful species, described from Hohenacker’s collection 
from the Kirovabad Region; the species is not represented in the 
Herbarium of the Botanical Institute of the Academy of Sciences of 
the USSR. Beck, having seen a type of this species, assigns it to the 
group Glandulosae G. Beck, which in our opinion, is extremely 
doubtful. No one has reported this species since Hohenacker. 


58. O. alba Steph. in Willd. Sp. pl. III (1800) 350; M.B. FI. taur.- 
cauc. II, 82; Reut. in DC. Prodr. XI, 20; Ldb. Fl. Ross. III, 316; Boiss. 
Fl. or. IV, 507; G. Beck, Monogr. Orob. 208, p.p. tab. IV, f. 64 and 
in Pflanzenr. IV, 261, 145, p.p. f. 14 (A—H); Schmalh. Fl. I, 295; 
Fedch. and Fler. Fl. 893, fig. 868; Grossh. Fl. Kavk. IV, 13, p.p.—0O. 
epithymum DC. FI. frang. III (1805) 490; Reut. l.c. 21; Ldb. I.c. 318; 
Boiss. l.c. 509, p.p.—O. glabrata C.A.M. in Eichw. Pl. casp.-cauc. I 
(1831) 16; Ldb. I.c. 322; ? Boiss. l.c. 511.—O. rubiginosa Dietrich, 
Fl. boruss. IV (1836) tab. 252.—? O. ferruginea C. Koch in Linnaea, 
XVII (1843) 291.—? O. psilandra C. Koch in Linnaea, XXII (1849) 
668.—O. hians Steven in Bull. Soc. Nat. Mosc. XXX (1857) 354.— 
O. wiedemanii Boiss. |.c. 510.—O. psilostemon Novopokr. in Bot. 
mat. Gerb. Bot. inst. AN SSSR, XIII (1950) 324.—Ic.: Rchb. I.c. Fl. 
Germ. XX, tab. 215, f. 1 and tab. 163; Dietrich, l.c. tab. 251 and 252; 
G. Beck, I.c.; Fedch. and Fler. |.c.; Hegi, Ill. Fl. VI, tab. 245, f. 1 and 
88 (a—d). 

Perennial. Plant up to 40 cm tall, glandular-pubescent (pubes- 
cence brownish or yellowish). Stem yellowish brown or reddish, mid- 
dle part 2-12 mm thick, base slightly thicker; covered with lanceolate 
or narrowly lanceolate, 15-22 mm long, scales. Inflorescence cylin- 
drical or oval, often lax and few-flowered, usually shorter than re- 
maining part of stem or equaling it. Bract scales ovate-lanceolate, 12— 
25 mm long, sometimes exceeding flower. Calyx segments free, 8-18 
mm long, entire, from ovate at base to narrowly lanceolate, often 
shortly subulate at tip, extremely rarely bidentate, light brown, with 
distinct darker veins, sometimes with reddish tinge. Corolla 15-25 
mm long, broadly campanulate, white, rarely light yellow, often with 
red or violet tinge in limb, glandular-pubescent outside, almost al- 
ways with mixture of colored (red or violet) glandular hairs; its lon- 
gitudinal dorsal line almost straight in middle, concave at base and at 
limb; upper lip sinuate, sometimes subentire, with extremely broad, 


94 


89 


orbicular lobes. Filaments more or less hairy for 1/3—1/4 length from 
base, glabrous above, but sparsely glandular-pubescent below anthers, 
rarely wholly glabrous, inserted 1-3 mm above base of corolla tube; 
anthers subglabrous. Style sparsely glandular-puberulent; stigma usu- 
ally red. May to July (Plate V, fig. 8). 

Parasitic on labiates, predominantly on species of genera Salvia 
L. and Thymus L.; in southern Ossetia on St. John’s-wort (Hypericum 
L.). Steppe, rubbly slopes, scrubs, subalpine meadows.— European 
USSR: Upper Dniester, Upper Dnieper, Volga-Don, Middle Dnieper, 
Bessarabia, Black Sea Region, Lower Don, Crimea. Caucasus: All 
regions. Soviet Central Asia: mountainous Turkmenia. General 
distribution: Central and Atlantic Europe, Scandinavia (southern part), 
western Mediterranean, Balkans-Asia Minor, Armenia-Kurdistan, Iran, 
India-Himalayas (?), Tibet (?). Described from Caucasus. Type in 
Leningrad. 

Note. A polymorphic species, highly variable in size and corolla 
color, length of bract scales, pubescence and other features. Beck 
distinguished a large number of forms and several varieties. The 
following may be noted: var. glabrata (C.A.M.) G. Beck [in Pflanzenr. 
IV, 261 (1930) 154] with subglabrous filaments, which is common in 
Transcaucasia, and var. bidentata G. Beck (1.c. 155) with bidentate 
calyx segments, rarely occurring in the Caucasus. Both these variants 
probably are independent physiological races. O. alba var. wiedemanii 
(Boiss.) G. Beck [Monogr. Orob. (1890) 212], as well as O. psilostemon 
Novopokr., probably are identical to var. glabrata (C.A.M.) G. Beck. 


59. O. pallidiflora Wimm. and Grab. FI. siles. II, 1 (1829) 233; 
Reut. in DC. Prodr. XI, 22; Schmalh. Fl. II, 296.—O. procera W. 
Koch, Fl.. Deutschl. IV (1833) 438; Reut. l.c. 16.—0O. bracteata 
Weinm. in Bull. Soc. Nat. Mosc. VII (1837) 65, non Viviani, 1830; 
Ldb. Fl. Ross. III, 317.—O. weinmanni Steud. Nomencl. bot. ed. 2, II 
(1841) 231.—O. antirrhina Reut. |.c. 21; Ldb. l.c. 318.—O. reticulata 
var. pallidiflora G. Beck, Monogr. Orob. (1890) 218, tab. IV, f. 67 (1) 
and in Pflanzenr. IV, 261 (1930) 159.—O. reticulata var. procera G. 
Beck, I.c. (1890) 218 and l.c. (1930) 160.—O. reticulata auct. non 
Wallr.; Fedch. and Fler. Fl. 894, fig. 869; Grossh. Fl. Kavk. IV, 13, 
p.p.—lIc.: Rchb. Ic. Fl. Germ. XX, tab. 166 and 168; G. Beck, l.c. 
(1890); Fedch. and Fler. lI.c. 

Biennial. Plant up to 60 cm tall, glandular-pubescent. Stem 
yellowish or brownish, sometimes tinged red, middle part 3-10 mm 
thick, base slightly thickened, covered with lanceolate, up to 20 cm 
long, scales. Inflorescence short cylindrical, fairly dense in upper part, 
lax below, many-flowered, shorter than or equaling remaining part of 


95 


90 


stem. Bract scales ovate-lanceolate, 12-16 mm long. Calyx segments 
free, 7-12 mm long, entire, from ovate base to narrowly lanceolate, 
short, subulate at tip, thick and dark colored, often blackish when dry, 
with less distinct veins. Corolla 15-20 mm long, campanulate, whitish 
or light yellow, limb often light violet or reddish colored, glandular- 
puberulent outside, hairs usually with mixture of colors (red or violet), 
longitudinal dorsal line of corolla straight in middle part, concave 
toward base and limb; upper lip sinuate, with broad, orbicular lobes. 
Filaments subglabrous, with few hairs at base, inserted 2-4 mm above 
base of corolla tube; anthers and style subglabrous.:Stigma usually 
red. June to July. 

Parasitic on species of genus Cirsium Adams.: C. oleraceum (L.) 
Scop., C. palustre (L.) Scop., C. heterophyllum (L.) Hill., C. arvense 
(L.) Scop., etc.; less commonly on other members of Compositae. 
Meadows, forest glades, scrub.—European USSR: Baltic Region, 
Ladoga-IImen, Volga-Kama, Upper Dniester, Middle Dniester, Volga- 
Don, Bessarabia, Black Sea Region, Lower Don, Crimea (?). Caucasus: 
Ciscaucasia. Western Siberia: Irtysh, Altai. General distribution: 
Scandinavia, Central and Atlantic Europe, western Mediterranean, 
Balkans. Described from Poland. Type in Kiel. 


60. O. reticulata Wallr. Orob. gen. diask. (1825) 42, non Reuter, 
1847; G. Beck, Monogr. Orob. 217, p.p. tab. IV, f. 67 (2) and in 
Pfianzenr. IV, 261, 155, p.p. f. 14 (J); Grossh. fl. Kavk. IV, 13, p.p.— 
O. scabiosae W. Koch, Deutschl. Fl. IV (1833) 440; Reut. in DC. 
Prodr. XI, 22.—Ic.: G. Beck, l.c. 

Biennial or perennial. Plant up to 25 cm tall, glandular-pubescent. 
Stem reddish, middle part 2—5 cm thick, base slightly thickened, with 
relatively few lanceolate, up to 10-15 mm long, scales. Inflorescence 
oblong or oval, usually fairly dense, shorter than remaining part of 
stem. Bract scales ovate-lanceolate or lanceolate up to 16 mm long. 
Calyx segments free, up to 10-12 mm long, entire, from ovate base 
narrowly lanceolate, short subulate at tip, thick and dark colored, usually 
blackish when dry, with indistinct veins. Corolla 15-20 mm long, 
campanulate, light yellow, limb with dark red or reddish violet tinge, 
often dark red throughout, glandular-pubescent outside, glandular-hairs 
of mixed colors over almost the entire surface (in red or violet), 
longitudinal dorsal line of corolla almost straight in middle part, concave 
toward base and limb; upper lip sinuate. Filaments subglabrous, inserted 
2-3 mm above base of corolla tube; anthers and style glabrous or 
subglabrous. Stigma red. July to August. 

Parasitic usually on species of genus Knautia L. and Scabiosa L.; 
also reported on some members of Compositae. Subalpine meadows, 


96 


91 


rubbly slopes and scrub in upper mountain belts— European USSR: 
Upper Dniester (Carpathia). Caucasus: Ciscaucasia, Dagestan, eastern, 
western and southern Transcaucasia. General distribution: Central 
Europe, western Mediterranean, Balkans-Asia Minor, Armenia- 
Kurdistan, India-Himalayas (?). Described from France. Type in Paris. 


61. O. grigorjevii Novopokr. in Bot. mat. Gerb. Bot. inst. AN 
SSSR, XIII (1950) 326. 

Perennial. Plant 15—30 cm tall, glandular-pubescent. Stem brownish 
yellow when dry, slightly thickened at base; covered with lanceolate, 
up to 20 mm long, scales. Inflorescence short cylindrical or oval, 
fairly dense, but lax at base, sometimes few-flowered, equaling or 
somewhat shorter than remaining part of stem. Bract scales ovate- 
lanceolate, usually recurved to narrowly lanceolate end, up to 20 mm 
long, at tip; on dorsal side fairly densely pubescent with short, reddish 
colored, glandular hairs. Calyx segments up to 20 mm long, free, 
entire, ovate at base, lanceolate-subulate, brownish with darker but 
indistinct veins. Corolla 18-22 mm long, tubular-campanulate, 
yellowish brown when dry, often tinged red, glandular-pubescent 
outside, usually with mixed colored (red or violet) glandular hairs, its 
longitudinal dorsal line almost uniformly concavely arcuate; upper lip 
sinuate, with extremely broad, semi-orbicular lobes. Filaments sparsely 
pubescent only at base, inserted 3-4 mm above base of corolla tube; 
anthers and style subglabrous. July to September. 

Parasitic on species of Origanum L. Rubbly slopes, subalpine 
meadows.— Soviet Central Asia: Pamiro-Alai. General distribution: 
Iran (eastern part), Dzhungaria-Kashgaria (southern part). Described 
from eastern Tadzhikistan (Khingou River basin). Type in Leningrad. 


62. O. raddeana G. Beck, Monogr. Orob. (1890) 194, tab. III, f. 
58.—O. alba var. raddeana G. Beck in Pflanzenr. IV, 261 (1930) 
154; Grossh. Fl. Kavk. IV, 13.—Ic.: G. Beck, lI.c. (1890). 

Perennial. Plant up to 30 cm tall, glandular-pubescent. Stem 
yellowish brown, often with reddish tinge, slightly thickened at base; 
covered with broadly lanceolate, up to 20 mm long, scales. 
Inflorescence short cylindrical or oval, usually lax and relatively few- 
flowered, shorter than or equaling remaining part of stem. Bract scales 
ovate-lanceolate, usually recurved at top to narrowly lanceolate tip, 
up to 20 mm long. Calyx segments free, 8-16 mm long, from ovate 
base narrowly lanceolate, entire, light brown, often tinged red, with 
prominent, darker veins. Corolla 16—23 mm long, campanulate, often 
with hairs of mixed colors, its longitudinal dorsal line almost equally 
concavely arcuate; upper lip smoothly sinuate. Filaments densely hairy 


97 


92 


over almost entire length, inserted 2-3 mm above base of corolla tube; 
anthers subglabrous. Style densely glandular-pubescent; stigma usually 
dark red. June to August (Plate V, fig. 9). 

Parasitic on species of Campanula L., usually on C. collina M.B.; 
subalpine meadows, rubbly slopes of upper mountainous belt.— 
Caucasus: All regions besides Talysh. General distribution: Asia Minor, 
Armenia-Kurdistan, Iran-Afghanistan (western part). Described from 
Georgia. Type in Leningrad (?). 


Series 3. Speciosae G. Beck, Monogr. Orob. (1890) 224 and in 
Pflanzenr. IV, 261 (1930) 136.—Corolla usually campanulate, 15-30 
mm long, glandular-pubescent outside, glandular hairs colorless, its 
longitudinal dorsal line almost straight in middle part. Calyx segments 
entire or bidentate, usually more or less subulate at tip. Pubescence, 
at least in inflorescence, usually with a mixture of longer and stiff 
whitish hairs. 


63. O. transcaucasica Tzvel. sp. n. in Addenda XXII, 686. 

Perennial or biennial (?). Plant up to 40 cm tall, glandular- 
pubescent with yellowish hairs. Stem reddish, middle part up to 6 mm 
thick, base slightly thicker and densely covered with smaller scales, 
above sparsely covered with lanceolate scales up to 20 mm long. 
Inflorescence short cylindrical, fairly lax, usually considerably shorter 
than remaining part of stem. Bract scales broadly lanceolate, up to 20 
mm long, equaling or somewhat longer than corolla tube, brownish, 
pubescence shorter on dorsal side, accumbent, with yellowish hairs. 
Calyx segments free in front, connate to a greater part at back (from 
the sides of axis) usually with 5 shorter teeth, light brown, with less 
prominent veins, lateral teeth linear-lanceolate, often dissimilar, 
equaling or longer than connate part of segments. Corolla 18-22 mm 
long, tubular-campanulate, limb pinkish violet, with darker veins, light 
brown or whitish (?) toward base, sparsely glandular-pubescent outside, 
longitudinal dorsal line of corolla almost straight in middle part, 
concave toward base and limb; upper lip weakly sinuate, with broad, 
semi-orbicular, lobes. Filaments hairy for 1/3 of length from base, 
glabrous above, but glandular-pubescent below anthers, inserted 1.5— 
2 mm above base of corolla tube; anthers glabrous. Style sparsely 
glandular-puberulent; stigma reddish. May to June. 

Host plant not determined. Forests, scrub.— Caucasus: Eastern 
Transcaucasia. Endemic. Described from Shemakha Region (environs 
of the village of Sharadil in hornbeam-oak forest at nearly 800 m 
above sea level; 6.VI.1941; collector unknown). Type in Baku. 


98 


93 


64. O. owerinii G. Beck in Fedde, Repert. XVIII (1922) 39 and 
in Pflanzenr. IV, 261 (1930) 140; Grossh. Fl. Kavk. IV, 12.—0O. 
crenata var. owerinii G. Beck, Monogr. Orob. (1890) 227. 

Perennial or biennial. Plant 15-60 cm tall, glandular-pubescent, 
inflorescence with mixture of longer, stiff whitish hairs. Stem reddish 
brown when dry, slightly thicker at base, with relatively few lanceolate, 
15-25 cm long, scales. Inflorescence short cylindrical or oval, lax, 
often few-flowered, shorter than or equaling remaining part of stem. 
Bract scales lanceolate, 15-25 mm long, with whitish pubescence on 
dorsal side. Calyx segments free, entire or rarely bidentate, up to 15 
mm long, at tip subulate, usually shorter than corolla tube. Corolla 
15-30 mm long, campanulate, yellowish at base, more or less violet- 
purple toward limb; color usually partially persists when dry, densely 
glandular-hairy outside, longitudinal dorsal line of corolla almost 
straight in middle part. Filaments hairy up to 1/3 to 1/2 of length from 
base, glabrous above, but often glandular-pubescent below anthers, 
inserted 2-3 mm above base of corolla tube. Anthers and style 
subglabrous. June to August. 

Parasitic on legumes, usually on species of Trifolium L., Lathyrus 
L. and Vicia L. Subalpine meadows, scrub, forests.— Caucasus: All 
regions. General distribution: Armenia-Kurdistan Described from 
Georgia (environs of Tbilisi). Type in Leningrad. 

Note. A polymorphic species, represented by _ several 
morphologically weakly differentiated physiological races, one of 
which, apparently, is var. woronowii G. Beck [in Fedde. Repert. XVIII 
(1922) 39] with 15-20 mm-long flowers. The most typical form is 
parasitic on Trifolium canescens Willd. It is closely related to the 
following species and replaces it in the upper montane zone. 


65. O. crenata Forsk. Fl. Aeg. Arab. (1775) 113; Reut. in DC. 
Prodr. XI, 37; G. Beck, Monogr. Orob. 225; p.p. tab. IV, f. 69 and in 
Pflanzenr. IV, 261, 136, p.p. f. 14 (L); Grossh. Fl. Kavk. IV, 12.—0O. 
speciosa DC. FI. frang. VI (1815) 393; Reut. l.c. 19; Boiss. Fl. or. IV, 
506.—O. pruinosa Lapeyr. Abreg. des pl. Pyren. Suppl. (1818) 87; 
Ldb. FI. Ross. III, 316.—Ic.: Rchb. Ic. Fl. Germ. XX, tab. 161; G. 
Beck, l.c. 

Perennial, biennial or annual. Plant up to 50 cm tall, glandular- 
pubescent, with mixture of longer, stiff, whitish hairs at least, in 
inflorescence. Stem yellowish or brownish, slightly thickened at base, 
covered with lanceolate, up to 25 mm long, scales. Inflorescence 
cylindrical or oblong, fairly dense or lax in lower part, many-flowered, 
shorter than or equaling remaining part of stem. Bract scales broadly 
lanceolate or lanceolate, 15-25 mm long, usually densely whitish hairy 


10 


— 


94 


on dorsal side. Calyx segments free, 10-20 mm long, bidentate or 
entire, subulate at tip usually almost equaling corolla tube. Corolla 
20-30 mm long, campanulate or tubular-campanulate, light yellow or 
whitish, sometimes with pale pinkish or violet tinge in limb, discolored 
on drying, glandular-pubescent outside, its longitudinal dorsal line 
almost erect in middle; upper lip smoothly sinuate, with broad, semi- 
orbicular lobes. Filaments usually hairy for no longer than 1/3 the 
length from base, inserted 2-3 mm above base of corolla tube. Anthers 
and style subglabrous. May to July. 

Parasitic on legumes; host plant not precisely docemimed Rubbly 
slopes, scrub, forests.— European USSR: Crimea (southern part). 
Caucasus: All regions, besides Talysh. General distribution: 
Mediterranean, Balkans-Asia Minor, Armenia-Kurdistan, Iran- 
Afghanistan (western part). Described from Egypt. Type in Kiel. 

Note. Just like the foregoing one, this species is polymorphic one, 
highly variable in the size of the corolla, shape of calyx segments, 
pubescence and other characters. When more herbarium material is 
available, probably some of the physiological races can be treated as 
distinct species. Not always clearly distinguished from the foregoing 
species and from O. picridis F. Schultz. 

Economic importance. In western Europe it is sometimes harmful 
to peas, beans, lentils, pea-vine; in the USSR it has so far not been 
observed on cultivated plants. 


Series 4. Minores G. Beck in Hal. and Braun, Nachr. Fl. Nied.- 
Oesterr. (1882) 135; Monogr. Orob. 136 and in Pflanzenr. IV, 261, 
167.—Corolla tubular-campanulate, 10-20 mm long, with colorless 
glandular hairs outside, its longitudinal dorsal line almost straight in 
middle part. Calyx segments entire or bidentate, tip usually subulate. 
Pubescence usually with mixture of long, stiff, whitish hairs, rarely 
without these. 


66. O. picridis F. Schultz. in Flora (1830) 500, sine descr. and in 
W. Koch, Fl. Deutsch]. IV (1833) 453; Reut. in DC. Prodr. XI, 26; G. 
Beck, Monogr. Orob. 245, tab. IV, f. 79 and in Pflanzenr. IV, 261, 
200; Grossh. Fl. Kavk. IV, 12.—0O. loricata auct. non Rchb.; Ldb. FI. 
Ross. III, 320, p.p.—Ic.: Rchb. Ic. Fl. Germ. XX, tab. 175 (a); G 
Beck, Monogr. Orob.; Hegi, Ill. Fl. VI, f. 89 (g—h). 

Biennial or perennial. Plant up to 40—50 cm tall, densely pubescent 
upwards with stiff whitish hairs, rarely only a mixture with shorter 
glandular pubescence. Stem yellowish, often tinged red, slightly 
thickened at base, covered with lanceolate, up to 22 mm long, scales. 
Inflorescence cylindrical, many-flowered, dense in upper part, lax in 


99 


; Plate VI. 
1. Orobanche gracilis Smith, flower. 2. O. colorata C. Koch, general appearance 
of plant, flower. 3. Boschniakia rossica (Cham. and Schlechtend.) B. Fedtsch., 
general appearance of plant, flower. 4. Mannagettaea ircutensis M. Pop. 5. 
Phacellanthus tybiflorus Sieb. and Zucc., general appearance of plant, flower. 


102 


96 


lower, equaling remaining part of stem, sometimes longer or somewhat 
shorter. Bract scales ovate-lanceolate, dorsal side fairly densely whitish 
hairy, 8-20 mm long. Calyx segments free, up to 15 mm long, bidentate 
or entire, subulate at tip, often equaling corolla tube. Corolla 16-20 
mm long, tubular-campanulate, light yellow, along veins sometimes 
violet or reddish colored, glandular-pubescent outside, its longitudinal 
dorsal line almost straight in middle part. Filaments usually densely 
hairy for almost half length from base, above glabrous or subglabrous, 
inserted 2-3 mm above base of corolla tube. Anthers and style 
subglabrous. May to July. 

Parasitic on members of Compositae, predominantly on species of 
genus Picris L. Steppe, rubbly slopes, scrub, fields— European USSR: 
Upper Dniester, Lower Don, Lower Volga. Caucasus: All regions. 
General distribution: Scandinavia, Central and Atlantic Europe, 
Mediterranean, Balkans-Asia Minor, Armenia-Kurdistan (?). Described 
from Germany. Type in Berlin (?). 

Note. Occupies a position intermediate between the groups 
Speciosae G. Beck and Minores G. Beck. A species closely resembling 
this may be found in the extreme western European part of USSR.— 
O. loricata Rchb. pat. [Iconogr. VII (1829) 41, f. 917], differing in 
the filaments, glandular-pubescent in the upper part, and in being 
parasitic on Artemisia campestris L. 


67. O. minor Sutt. in Trans. Linn. Soc. IV (1797) 179; Reut. in 
DC. Prodr. XI, 29, p.p. Ldb. Fl. Ross. III, 320; Boiss. Fl. or. IV, 512; 
G. Beck, Monogr. Orob. 251, tab. IV, f. 82 and in Pflanzenr. IV, 261, 
205, f. 15 (A—G); Fedch. and Fler. Fl. 895, fig. 872; Grossh. Fl. Kavk. 
IV, 12.—? O. barbata Poir. in Lam. Encycl. IV (1796) 621.—0O. 
concolor Duby, Bot. Gall. I (1828) 350; Reut. I.c. 231; ? Ldb. l.c. 
318; G. Beck, Monogr. Orob. 256.—Ie.: Rchb. Ic. Fl. Germ. XX, tab. 
183; G. Beck, I.c.; Fedch. and Fler. |.c.; Hegi, Ill. Fl. VI, f. 90 (a-e). 

Biennial or perennial. Plant 10-35 cm tall, glandular-pubescent, 
usually with mixture of longer whitish hairs in inflorescence. Stem 
reddish brown, brownish or yellowish, middle part 1.5-4 mm thick, 
base sometimes clavate, with relatively few lanceolate, up to 20 mm 
long, scales. Inflorescence cylindrical of oblong, lax, shorter than or 
equaling remaining part or stem. Bract scales ovate-lanceolate, 7-12 
mm long, dorsal side fairly densely white hairy. Calyx segments free, 
4-12 mm long, incised for half or more of length into 2 lanceolate- 
subulate teeth, rarely entire. Corolla 10-18 mm long, tubular- 
campanulate, light yellow, usually with violet tinge in limb and along 
veins, sparsely glandular-pubescent outside, its longitudinal dorsal line 
almost straight in middle. Filaments sparsely hairy, glabrous above, 


103 


97 


inserted 3—5 mm above base of corolla tube, usually somewhat below 
the middle of tube. Anthers and style subglabrous. Stigma usually red, 
rarely yellow. May to June. 

Parasitic on legumes, predominantly on species of Trifolium L. 
Rubbly slopes, scrub.— European USSR: Bessarabia, Crimea (southern 
part). Caucasus: Western Transcaucasia, Talysh. General distribution: 
Scandinavia (southern part), Central and Atlantic Europe, 
Mediterranean, Balkans-Asia Minor, Africa, introduced into Central 
America and New Zealand. Described from Great Britain. Type in 
London. 

Note. In the Adzharian ASSR, plants that are completely light 
yellow in color, with smaller flowers and highly reduced, entire calyx 
segments.—var. concolor (Duby) G. Beck [in Pflanzenr. IV, 261 (1930) 
214]—occasionally occur together with typical specimens. This variety, 
which evidently represents a special physiological race, probably was 
erroneously reported from Volhynia by Ledebour (Flora rossica). 

Economic importance. In western Europe it sometimes harms 
clover fields. 


68. O. versicolor F. Schultz in Flora, I (1843) 129; G. Beck, 
Monogr. Orob. 237, tab. IV, f. 75 and in Pflanzenr. IV, 261, 183; 
Fedch. and Fler. Fl. 894, fig. 871; Grossh. Fl. Kavk. IV, 12.—0. 
pubescens Urv. Enum. pl. or. (1822) 76, No. 567, non Gilib. 1781; 
Reut. in DC. Prodr. XI, 27; Boiss. Fl. or. IV, 507.—O. arachnoidea 
F. Schultz in Linnaea, XIX (1845) 26 (ex diagn.) and apud G. Koch 
in Linnaea, XXII (1849) 669; Schmalh. Fl. II, 298.—O. squalida 
Steven in Bull. Soc. Nat. Mosc. XXX (1857) 354; Schmalh. FI. II, 
298.—Ic.: G. Beck, l.c.; Fedch. and Fler. l.c. 

Biennial, annual or perennial (?). Plant up to 30 cm tall, glandular- 
pubescent, usually with mixture of longer whitish hairs. Stem yellowish 
or brownish, sometime tinged red, middle part 2-7 mm thick, base 
more or less thickened, covered with lanceolate, acute, 8-20 mm long, 
scales. Inflorescence short cylindrical or oblong, fairly dense or lax, 
many-flowered, usually shorter than remaining part of stem. Bract 
scales ovate-lanceolate, gradually acuminate, 8-15 mm long, dorsal 
side usually whitish hairy. Calyx segments free, 7-12 mm long, 
bidentate or rarely entire, their teeth often dissimilar, lanceolate- 
subulate. Corolla 10-15 mm long, tubular-campanulate, light yellow, 
often with lilac or violet tinge, sometimes brownish in limb, glandular- 
pubescent outside, but above, especially close to throat, with mixture 
of long, tangled, often whitish lanate hairs; its longitudinal dorsal line 
nearly straight in middle. Filaments densely hairy close to base, 
sometimes hairy almost to half length, above glabrous, inserted 3-5 


104 


98 


mm above base of corolla tube, usually somewhat lower than middle 
of tube. Anthers and style glabrous or subglabrous. May to June. 

According to Beck, parasitic on umbellifers and members of 
Compositae as well as legumes; in Crimea also observed on annual 
umbellifers of genera Torilis Adams, Orlaya Hoffm., Caucalis L., etc. 
Rubbly slopes, scrubs.—European USSR: Crimea (southern part). 
Caucasus: Western Transcaucasia. General distribution: Mediterranean, 
Balkans-Asia Minor, Armenia-Kurdistan. Described from Greece (?) 
Type in Paris (?). 


69. O. hederae Duby, Bot. Gall. I (1828) 350; Reut. in DC. Prodr. 
XI, 28; Boiss. Fl. or. IV, 513; G. Beck, Monogr. Orob. 259, tab. IV, 
fig. 83 and in Pflanzenr. IV, 261, 219, fig. 15 (H); Fedch. and Fler. 
Fl. 895, fig. 873; Grossh. Fl. Kavk. IV, 12.—Ic.: Rchb. Ic. Fl. Germ. 
XX, tab. 182; G. Beck, l.c.; Fedch. and Fler. |.c.; Hegi, Ill. Fl. VI, fig. 
90 (f-h). 

Perennial. Plant up to 35 cm tall, glandular-pubescent, often with 
mixture of longer whitish hairs. Stem yellowish or reddish, middle 
part 3-10 mm thick, usually clavate or tuberous at base, covered with 
lanceolate or ovate-lanceolate, 10-30 mm long, scales. Inflorescence 
usually cylindrical, many-flowered, upper part fairly dense, lower lax, 
more or less equaling or longer than remaining part of stem. Bract 
scales ovate-lanceolate or lanceolate, 10-18 mm long. Calyx segments 
free, 7-14 mm long, entire or bidentate, lanceolate-subulate at tip. 
Corolla 14-18 mm long, broadly triangular, yellowish, whitish toward 
base, sparsely glandular-puberulent or subglabrous outside, tube 
distinctly constricted below throat, somewhat swollen closer to base, 
longitudinal dorsal line nearly straight in middle. Filaments sparsely 
pubescent at base, glabrous above, inserted 3-5 mm above base of 
corolla tube, usually somewhat below its middle. Anthers and style 
glabrous or subglabrous. May to July (Plate V, fig. 10). 

Parasitic on species of ivy (Hedera L.). Forests, scrub, parks.— 
European USSR: Crimea (southern part). Caucasus: All regions, besides 
southern Transcaucasia. General distribution: Central and Atlantic 
Europe, Mediteranean, Balkan States-Asia Minor. Described from 
France. Type in Berne. 


70. O. connata C. Koch in Linnaea, XXII (1849) 660; Boiss. FI. 
or. IV, 504; G. Beck, Monogr. Orob. 262 and in Pflanzenr. IV, 261, 
218; Grossh. Fl. Kavk. IV, 12. 

Biennial or perennial (?). Plant about 25 cm tall, glandular- 
pubescent. Stem subglabrous, with relatively few lanceolate scales. 
Inflorescence cylindrical, lax. Bract scales lanceolate, dorsal side 


ee 


105 


99 


glandular-pubescent. Calyx segments connate in front and at back, 
broadly oval, bidentate, considerably shorter than corolla tube, their 
teeth from deltoid base to short-acuminate, shorter than calyx tube. 
Corolla 15—18 mm long, tubular, brownish-red when dry, glandular- 
pubescent outside, its longitudinal dorsal line almost straight in middle 
part. Filaments inserted 4-S mm above base of corolla tube, hairy up 
to almost half length from base, glabrous above. Anthers and style 
subglabrous. May to June. 

Host plant not determined. Rubbly slopes.— Caucasus: Southern 
Transcaucasia. General distribution: Armenia-Kurdistan (?). Described 
from Armenia (environs of Yerevan). Type was in Berlin. 

Note. A doubtful species, so far known only from the authentic 
specimens of Koch. Following Beck, who has seen these specimens, 
we refer this species to the group Minores G. Beck. 


Series 5. Curvatae G. Beck in Hal. und Braun, Nachtr. Fl. Nied.- 
Oesterr. (1882) 128; Monogr. Orob. 134 and in Pflanzenr. IV, 261, 
243.—Corolla tubular-campanulate or broadly tubular, 14—25 mm long, 
with colorless, glandular hairs outside, usually in different shades of 
yellow, its longitudinal dorsal line more or less equally concave over 
its entire length. Calyx segments bidentate or entire. Pubescence usually 
brownish or yellowish, rarely with a mixture of whitish hairs. 


71. O. armena Tzvel. sp. n. in Addenda, XXII, 686. 

Biennial or perennial (?). Plant up to 35-40 cm tall, glandular- 
pubescent. Stem brownish-yellow, subglabrous below, somewhat 
thickened at base, covered with lanceolate, up to 25 mm long, scales. 
Inflorescence short cylindrical, upper part dense, lower lax, usually 
somewhat shorter than remaining part of stem. Bract scales broadly 
lanceolate, brownish, up to 25 mm long, sparsely glandular-pubescent, 
equaling corolla tube or somewhat longer. Calyx segments up to 15 
mm long, highly connate from the side of axis of inflorescence (for 
3-4 mm ) and, here, usually with 5 short teeth, front part free, light 
brown, with indistinct veins; their lateral teeth lanceolate, usually 
unequal, almost equaling connate parts of segments. Corolla 18-22 
mm long, tubular-campanulate, brownish yellow when dry, sparsely 
glandular-pubescent outside, its longitudinal dorsal line concave, not 
so strongly in lower half of corolla, upper lip slightly sinuate, with 
broad semi-orbicular lobes. Filaments densely hairy for 2/3 length 
from base, sparsely glandular-puberulent above, inserted 3-4 mm above 
base of corolla tube. Anthers pubescent along sutures of thecae. Style 
glandular-puberulent; stigma brownish. June. 


106 


100 


Host plant not determined. Rubbly slopes.— Caucasus: probably 
southern Transcaucasia. General distribution: Armenia-Kurdistan. 
Described from Turkey, close to its border with Armenia (Kagyzman, 
along Kagyzman-Darasu gully close to Araks, 18.VI.1886, V. 
Massal’skii). Type in Leningrad. 


72. O. kurdica Boiss. and Hausskn. in Boiss. Fl. or. IV, 505; G. 
Beck. Monogr. Orob. 161, tab. III, f. 52 and in Pflanzenr. IV, 261, 
260; ? Grossh. Fl. Kavk. IV, 13.—Ic.: G. Beck, Monogr. Orob. 

Biennial or perennial. Plant up to 50 cm tall, glandular-pubescent. 
Stem yellowish, middle part up to 12 mm thick, base slightly thicker; 
covered with fairly numerous, lanceolate, up to 30 mm long, scales. 
Inflorescence cylindrical, rarely lanceolate, lax, usually almost equaling 
remaining part of stem or longer. Bract scales lanceolate, up to 30 mm 
long, often equaling corolla tube, dorsal side more or less glandular- 
pubescent. Calyx segments free, up to 22 mm long, greatly varying in 
size and shape, incised usually up to more than half length into 2 
lanceolate light brown teeth. Corolla 20-25 mm long, tubular- 
campanulate, usually brownish yellow when dry, lighter toward base, 
sparsely glandular-pubescent outside, its longitudinal dorsal line mildly 
concave. Filaments pubescent almost from base to middle, glabrous 
above, but glandular-puberulent below anthers; inserted 3-4 mm above 
base of corolla tube; anthers glabrous or subglabrous. Style sparsely 
glandular-puberulent. June to July. 

Host plant not determined; reported on species of genera Tanacetum 
L., Cardus L., and Phlomis L. Rubbly, predominantly calcareous 
slopes.— Caucasus: Southern Transcaucasia. General distribution: Asia 
Minor, Armenia-Kurdistan. Described from Turkey. Type in Geneva, 
isotype in Leningrad. 

Note. A variant of this species with very large calyx segments and 
a subglabrous stem has been described from Mount Ararat beyond the 
boundaries of the Soviet Union. Beck has named it O. kurdica var. 
macrocalyx G. Beck (in Pflanzenr. IV (1930) 260), and it probably 
represents a distinct species. The specimen identified by Beck as “O. 
kurdica” from Abkhazia (probably, a slip of the pen), undoubtedly 
refers to O. colorata C. Koch and not to O. kurdica Boiss. and Hausskn. 


73. O. grossheimii Novopokr. in Bot. zhurn. SSSR, XXXIV, No. 
3 (1949) 282. 

Perennial. Plant 25—30 cm tall, relatively densely glandular- 
pubescent, often short rusty tomentose in inflorescence. Stem brownish, 
sometimes tinged red, middle part up to 15 mm thick, base somewhat 
thicker, covered with numerous lanceolate, up to 30 mm long, scales, 


107 


101 


often imbricate in lower part of stem and smaller. Inflorescence 
cylindrical, dense in upper part, lax below, equaling remaining part of 
stem or somewhat longer. Bract scales narrowly lanceolate, up to 30 
mm long, equaling corolla or longer, densely pubescent, often rusty 
tomentose on dorsal side. Calyx segments free or connate in front, up 
to 15 mm long, usually broad and bidentate, with lanceolate teeth, 
rarely narrower and entire. Corolla 20-25 mm long, tubular- 
campanulate, yellowish or yellowish brown, glandular-pubescent 
outside, its longitudinal dorsal line mildly, but more or less evenly, 
concave. Filaments densely hairy from base to 2/3 length, sparsely 
glandular-puberulent above, inserted 2-3 mm above base of corolla 
tube; anthers subglabrous. Style sparsely glandular-puberulent. June 
to August. 

Host plant not determined; probably (according to I.V. 
Novopokrovsky) it is Cephalaria gigantea (Ldb.) Bobr. Subalpine 
meadows, stony slopes.— Caucasus: Ciscaucasia, western, southern and 
eastern Transcaucasia. General distribution: probably Armenia- 
Kurdistan. Described from South Ossetia. Type in Leningrad. 

Note. I.V. Novopokrovsky considers this species to be close to O. 
vulgaris Poir.; however, in our opinion, it belongs to the group 
Curvatae G. Beck and closely resembles O. kurdica Boiss. and 
Hausskn. The possibility that this species belongs to one of the 
following species described by C. Koch cannot be ruled out: O. 
ferruginea C. Koch or O. quadrifida C. Koch. 


74. O. major L. Sp. pl. (1753) 632; G. Beck, Monogr. Orob. 169, 
tab. III, f. 45 and in Pflanzenr. IV, 261, 246; Fedch. and Fler. Fl. 892, 
fig. 864; Grossh. Fl. Kavk. IV.—O. elatior Sutt. in Transact. Linn. 
Soc., IV (1797) 178, tab. 17, f. 4; Reut. in DC. Prodr. XI, 25; Ldb. 
Fl. Ross. III, 319.—O. kochii F. Schultz. in Flora (1847) 66, tab. II, 
f. 1; Schmalh. Fl. II, 297.—Ic.: Sutt. l.c.; Rchb. Ic. Fl. Germ. XX tab. 
173; F. Schultz, l.c.; Fedch. and Fler. l.c.; Hegi, Ill. Fl. VI, f. 84 (e— 
f).—Exs.: Kar. and Kir. Pl. exs. (1841) No. 1802 (f. karelinii G. 
Beck); Kerner, Fl. exs. austro-hung. No. 913. 

Perennial or biennial. Plant 15-40 cm tall, glandular-pubescent 
(pubescence brownish). Stem reddish or brownish, middle part 3-8 
mm thick, base somewhat thicker, covered with fairly numerous, 
lanceolate, 10-20 mm long, scales. Inflorescence cylindrical, rarely 
oval, dense in upper part, lax below, few-flowered, usually almost 
equaling remaining part of stem. Bract scales broadly lanceolate, 12— 
18 mm long. Calyx segments free, brownish, fairly thick, 9-13 mm 
long, usually broad and bidentate, with narrow lanceolate teeth, rarely 
entire. Corolla 17-22 mm long, tubular-campanulate, pale yellow or 


108 


102 


whitish, often tinged pink, glandular-pubescent outside, its longitudinal 
dorsal line mildly, more or less evenly, concave. Filaments hairy for 
1/4-—1/3 of length from base, glabrous above, but glandular-puberulent 
below anthers, inserted 3-6 mm above base of corolla tube; anthers 
glabrous or subglabrous. Style glandular-puberulent. June to July. 

Parasitic on composites, usually on species of genera Centaurea 
L. and Echinops L. Steppe, rubbly slopes, scrub.— European USSR: 
Baltic Region, Upper Dniester, Upper Dnieper, Middle Dnieper, Volga- 
Don, Volga-Kama, Trans-Volga Region, Bessarabia, Black Sea Region, 
Lower Don, Lower Volga, Crimea. Caucasus: Ciscaucasia, Dagestan, 
western Transcaucasia (southern part), southern Transcaucasia, Talysh. 
Western Siberia: Upper Tobol, Talysh, Altai. Soviet Central Asia: 
Aralo-Caspian, Baltic Region, Dzhungaria-Tarbagatai, Tien Shan, Syr 
Darya, Pamiro-Alai. General distribution: Scandinavia, Central and 
Atlantic Europe, Western Mediterranean, Balkans-Asia Minor, Armenia- 
Kurdistan, Iran, India-Himalayas, Dzhungaria-Kashgaria. Described 
from Western Europe. Type in London. 

Note. A polymorphic species, containing several slightly 
morphologically different physiological races, not yet sufficiently 
investigated. One of the foregoing forms—f. karelinii G. Beck 
[Monogr. Orob. (1890) 171], with predominantly entire calyx segments, 
apparently, is usually found in Soviet Central Asia. In the Caucasus, 
this species becomes very rare, but reappears along the southern borders 
of the USSR representing here separate physiological races, often 
difficult to distinguish from O. kurdica Boiss. and Hausskn. 


75. O. flava Mart. in F. Schultz, Beitrag zur Kenntn. deutsch. 
Orob. (1829) 9; Reuter in DC. Prodr. XI, 27; G. Beck, Monogr. Orob. 
181, tab. III, f. 53 and in Pflanzenr. IV, 261; Fedch. and Fler. Fl. 893, 
fig. 866; Grossh. Opred. rast. Kavk. 256.—Ic.: Rchb. Ic. Fl. Germ. 
XX, tab. 177; G. Beck, I.c.; Fedch. and Fler. I.c.; Hegi, Ill. Fl. VI, tab. 
245, f. 86 (e-h). 

Perennial. Plant up to 60-70 cm tall, glandular-pubescent. Stem 
yellowish, brownish when dry, with relatively few early dying 
lanceolate, up to 20 mm long, scales. Inflorescence cylindrical or 
oblong, upper part fairly dense, lower lax, usually considerably shorter 
than or rarely equaling remaining part of stem. Bract scales broadly 
lanceolate, up to 20 mm long, dorsal side fairly densely grayish- 
pubescent. Calyx segments free, up to 15 mm long, entire or bidentate, 
short subulate at tip. Corolla 15-20 mm long, tubular-campanulate, 
yellow or light yellow, often brownish yellow when dry, glandular- 
pubescent outside, its longitudinal dorsal line evenly concave. Filaments 
densely hairy for its entire length from base, sparsely glandular- 


109 


103 


pubescent above, inserted 3-6 mm above base of corolla tube; anthers 
and style almost glabrous. June to August. 

Parasitic on composites, usually on species of genera Petasites 
Gaertn. and Tussilago L. Along banks of mountain rivulets and streams, 
scrub, forests.—European USSR: Upper Dniester (?). Caucasus: 
Ciscaucasia, Dagestan, western and eastern Transcaucasia. General 
distribution: Central Europe, western Mediterranean, Balkans. Described 
from Germany. Type in Miinich. 


76. O. inulae Novopokr. and Abram. in Bot. mat. Gerb. Bot. inst. 
AN SSSR, XIII (1950) 323 and in Grossh. Opred. rast. Kavk. (1949) 
730. 

Perennial. Plant up to 30 cm tall, glandular-pubescent. Stem yellow 
or light brown, with relatively few lanceolate, up to 15 mm long, 
scales. Inflorescence short cylindrical or oblong, upper part fairly dense, 
lower lax, shorter than or rarely equaling remaining part of stem. 
Bract scales ovate-lanceolate, up to 18 mm long. Calyx segments free, 
up to 15 mm long, entire, rarely bidentate, usually short subulate at 
tip. Corolla 15-20 mm long, tubular-campanulate, light yellow, usually 
brownish yellow when dry, glandular-pubescent outside, its longitudinal 
dorsal line almost evenly concave. Filaments hairy for almost its entire 
length from base, sparsely glandular-pubescent above, inserted 1.5-3 
mm above base of corolla tube. Anthers and style subglabrous. June 
to August (Plate V, fig. 11). 

Parasitic on species of genus Inula L. (I. grandiflora Willd., I. 
cordata Boiss., I. magnifica Lipsky, etc.). 

Subalpine meadows, scrub, rubbly slopes.— Caucasus: Ciscaucasia, 
Dagestan, western and eastern Transcaucasia. Endemic. Described from 
South Ossetia. Type in Leningrad. 


77. O. krylowii G. Beck in Kryl. Mater. k fl. Permsk. gub. II 
(1881) 202 (without diagn.) and in Oest. Bot. Zeitschr. (1881) 309; 
Kryl. Fl. Zap. Sib. X, 2544.—O. major f. krylowii G. Beck, Monogr. 
Orob. (1890) 171, tab. III, f. 45 (3) and in Pflanzenr. IV, 261, 246; 
Kryl, Fl. Alt. 1 Tomsk. gub. IV, 996.—Ic.: G. Beck, l.c. 

Perennial. Plant up to 40 cm tall, subglabrous below, sparsely 
appressed-pubescent above. Stem brownish yellow, usually tinged red, 
middle part 3-9 mm thick, somewhat thickened at base, covered more 
abundantly in lower part of stem with broadly lanceolate, 10-20 mm 
long, scales, glabrous. Inflorescence cylindrical or oblong, fairly dense 
or lax, usually shorter than remaining part of stem. Bract scales ovate 
or broadly lanceolate, 12-18 mm long, sparsely glandular-pubescent. 
Calyx segments free, 6-12 mm long, usually not exceeding middle of 


110 


104 


corolla tube; entire or unevenly bidentate, short-subulate at tip. Corolla 
17-22 mm long, broadly tubular, light yellow, densely glandular- 
pubescent outside, its longitudinal dorsal line almost evenly arcuate. 
Filaments densely hairy for 2/3 length from base, sparsely glandular- 
hairy above, inserted 3—4 mm above base of corolla tube; anthers 
pubescent along sutures of thecae. Style sparsely, glandular-pubescent. 
June to July. 

Parasitic on species of Thalictrum L. Forests, scrub.— European 
USSR: Volga-Kama. Western Siberia: Ob’ (southern part), Upper Tobol, 
Irtysh, Altai. Eastern Siberia: Angara-Sayan. Soviet Central Asia: Tien 
Shan, Dzhungaria-Tarbagatai, Balkhash region (eastern part). General 
distribution: Dzhungaria-Kashgar. Described from Perm Region (banks 
of Chusovaya River). Type in Leningrad. 


78. O. alsatica Kirschl. Prodr. fl. d’Alsace (1836) 109; G. Beck, 
Monogr. Orob. 186, tab. III, f. 49 and in Pflanzenr. IV, 261, 256, f. 
17 (G); Fedch. and Fler. Fl. 892, fig. 865; Grossh. Fl. Kavk. IV, 13.— 
O. cervariae Suard in Godr. Fl. Lorr. II (1843) 180; Schmalh. FI. II, 
296.—O. brachysepala F. Schultz, Arch. fl. France et Allem. (1844) 
808; Reut. in DC. Prodr. XI, 57.—O. libanotidis Rupr. in Beitr. zur 
Plfanzenr. Russ. Reich, IV (1845) 70; Ldb. Fl. Ross. III, 319; Schmalh. 
Fl. II, 296; Kryl. Fl. Zap. Sib. X, 2546.—O. major var. minor Meinsh. 
Fl. ingr. (1878) 263.—Ic.: Rchb. Ic. Fl. Germ. XX, tab. 174; G. Beck, 
Monogr. Orob.; Fedch. and Fler., l.c.; Hegi, Ill. Fl. VI, f. 85 and 86 
(a—d). 

Biennial or perennial. Plant up to 60 cm tall, glandular-pubescent. 
Stem yellow or yellowish brown, middle part 3-10 mm thick, base 
often clavate or tuberous, covered with fairly numerous, broadly 
lanceolate, 12-22 mm long, scales. Inflorescence cylindrical or oblong, 
dense or in lower part lax, with distant flowers, usually considerably 
shorter than remaining part of stem. Bract scales broadly lanceolate, 
10-20 mm long, dorsal side fairly densely, often whitish hairy. Calyx 
segments free or connate in front, 7-12 mm long, usually bidentate, 
with lanceolate-subulate teeth, rarely entire, light brown. Corolla 14— 
22 mm long, broadly tubular, yellowish, often with lilac tinge in limb, 
usually brownish yellow when dry, sparsely glandular-pubescent 
outside, its longitudinal dorsal line nearly evenly concave. Filaments 
hairy up to 1/3-1/4 length from base, rarely subglabrous or densely 
hairy for 2/3 length; anthers pubescent along sutures of thecae. Style 
glabrous or subglabrous. June to August. 

Parasitic on umbellifers; observed on species of genera Libanotis 
Cr., Peucedanum L., Heracleum L., Aegopodium L. etc. Steppe, rubbly 
slopes, scrub, forests, subalpine meadows.— European USSR: Baltic 


111 


105 


Region. Ladoga-IlImen, Volga-Kama, Upper Dniester, Upper Volga, 
Middle Dnieper. Volga-Don, Trans-Volga, Bessarabia, Black Sea 
Region, Lower Don, Lower Volga, Crimea. Caucasus: Ciscaucasia, 
western and southern Transcaucasia, Talysh. Western Siberia: Ob’ 
Region (southern part), Upper Tobol, Irtysh, Altai. Eastern Siberia: 
Angara-Sayan. Soviet Central Asia: Aralo-Caspian, Balkhash Region, 
Dzhungaria-Tarbagatai, Tien Shan. General distribution: Central and 
Atlantic Europe, western Mediterranean, Balkans-Asia Minor, Armenia- 
Kurdistan, Iran (?), Dzhungaria-Kashgaria. Described from France. Type 
in Strasbourg. 

Note. A polymorphic species. Usually, two varieties are 
distinguished: var. typica G. Beck [Monogr. Orob. (1890) 176] with 
flowers 18-22 mm long, and var. libanotibis (Rupr.) G. Beck (l.c. 
177) with flowers 14-18 mm long. Of these only the second occurs 
almost throughcut the range of the species, whereas the first is prevalent 
predominantly in the mountainous regions. These varieties, apparently 
are not physiological races (both occur on Libanotis Cr. and 
Peucedanum L.) and are not always clearly distinguishable. In the 
Caucasus, O. alsatica Kirschl: occurs rarely and not in all regions, but 
the polymorphism of this species is especially high here. Var. heraclei 
Tzvel. var. n. occurs in subalpine meadows of the Caucasian Range, 
where it is parasitic on species of Heracleum. It has a fairly lax 
inflorescence with a large corolla and is densely hairy over the entire 
length of the filaments. Var. iranica Tzvel. var. n. occur along the 
borders of Iran (where typical O. alsatica is absent) and also, probably, 
represents a physiological race with a large corolla and subglabrous 
filaments. 


79. O. rosea Tzvel. sp. n. in Addenda XXII, 687. 

Biennial or perennial (?). Plant up to 49 cm tall, densely glandular- 
pubescent. Stem yellowish brown, usually reddish to dark red above, 
middle part 6-10 mm thick, base slightly thicker; covered with fairly 
numerous, lanceolate up to 30 mm long, scales. Inflorescence 
cylindrical or oblong, fairly dense, but lax with distant flowers below, 
usually almost equaling or longer than remaining part of stem. Bract 
scales broadly lanceolae to linear lanceolate, 15-30 mm long, almost 
equaling corolla or longer, dorsal side densely glandular-pubescent. 
Calyx segments free, 10-20 mm long, usually bidentate, with narrowly 
lanceolate teeth, rarely entire, light brown. Corolla 16-22 mm long, 
broadly tubular, pinkish, light colored in limb, usually (although in 
part) preserves pinkish color of tube when dry, with brownish yellow 
limb, densely glandular-pubescent outside, its longitudinal dorsal line 
almost evenly concave. Filaments densely hairy in the lower half, 


112 


106 


sparsely glandular-pubescent above; anthers glabrous or subglabrous. 
Style sparsely glandular-pubescent. June to July. 

Parasitic on Prangnos ferulacea Lindl. Rubbly slopes, cliffs and 
crags from lower to upper mountain belts.—Caucasus: Southern 
Transcaucasia, Talysh. Soviet Central Asia: mountainous Turkmenia. 
General distribution: Armenia-Kurdistan, Iran. Described from 
Nakhichevan ASSR (peaks of low mountains 1.5—2 km east of city of 
Ordubad.—T. Egorova, N. Tzvelev and S. Czerepanov). Type in 
Leningrad. 

Note. Parasitic, like O. alsatica Kirschl., on umbellifers, this species 
also shows no less proximity to O. kurdica Boiss. et Hausskn. and O. 
grossheimii Novopokr. 


Series 6. C. cruentae G. Beck, Monogr. Orob. (1890) 135 and in 
Pflanzenr. IV, 261 (1930) 278.—Corolla colorless, tubular- 
campanulate, 16-22 mm long, hairs dark colored, yellow near base, 
red or brownish red in limb, inside with darker coloring and brilliance, 
its longitudinal dorsal line nearly evenly concave. Calyx segments 
usually bidentate, pubescence short, grayish or brownish. 


80. O. gracilis Smith in Transact. Linn. Soc. IV (1797) 172; G. 
Beck, Monogr. Orob. 195, tab. III, f. 59 and in Pflanzenr. IV, 261, 
279, f. 17 (A-C); Fedch. and Fler. Fl. 893, fig. 867; Grossh. Fl. Kavk. 
IV, 13.—O. cruenta Bertol. Rar. it. pl. dec. 3 (1810) 56; Reut. in DC. 
Prodr. XI, 15; Ldb. Fl. Ross. III, 315; Boiss. Fl. or. IV, 505.—Ic:: 
Rchb. Ic. Fl. Germ. XX, tab. 159; G. Beck, Monogr. Orob.; Fedch. 
and Fler. |.c.; Hegi, Ill. Fl. VI, tab. 245, f. 2. 

Perennial. Plant up to 40-45 cm tall, glandular-pubescent, 
sometimes subglabrous. Stem dark red or yellow with reddish tinge, 
middle part 2-6 mm thick, base more or less thickened, covered with 
lanceolate, up to 15 mm long, scales. Inflorescence cylindrical or 
oblong, fairly dense or lax in the lower part, usually almost equaling 
remaining part of stem. Bract scales ovate-lanceolate, 7-18 mm long, 
more or less whitish on dorsal side due to dense but short pubescence. 
Calyx segments free, usually incised up to middle into 2 narrowly 
lanceolate teeth, very rarely entire, 7-12 mm long. Corolla 16-22 mm 
long, tubular-campanulate or campanulate, yellow, dark red in upper 
part of limb, inner side usually bright colored and lustrous, sometimes 
very dark colored or yellow, sparsely glandular-puberulent outside, 
longitudinal dorsal line almost evenly concave. Filaments inserted 1- 
2 mm above base of corolla tube, hairy for 1/4 to 1/3 from base, 
glabrous above, but often glandular-pubescent below anthers; anthers 


113 


107 


subglabrous. Style sparsely glandular-puberulent. June to August (Plate 
VI, fig. 1). 

Parasitic on legumes; observed on species of genera Cytisus L., 
Genista L., Dorycnium L., Coronilla L., Argyrolobium Eckl. et Zeyh., 
etc. Rubbly slopes, scrub, subalpine meadows.— European USSR: 
Upper Dniester (Carpathian Mts.); Bessarabia(?). Caucasus: 
Ciscaucasia, western Transcaucasia, eastern Transcaucasia (western 
part). General distribution: Central Europe, western Mediterranean, 
Balkans-Asia Minor. Described from Italy. Type in London. 


Series 7. Arcuatae G. Beck, Monogr. Orob. (1890) 135. and in 
Pflanzenr. IV, 261, 269.—Corolla tubular-campanulate, 16-25 mm 
long, colorless, with dark colored hairs, brown or brownish-violet, 
more or less uniformly colored; upper lip almost entire, galeate, its 
longitudinal dorsal line in the upper half strongly, and lower half 
mildly, concave. Calyx segments usually bidentate. Pubescence usually 
short, brownish, sometimes white lanate, or entire plant subglabrous. 


81. O. colorata C. Koch in Linnaea, XVII (1843) 291; Ldb. FI. 
Ross. III, 322.—O. anatolica auct. fl. cauc. non Boiss. et Reut. 

Perennial. Plant up to 20-32 cm tall, glabrous or diffusely 
glandular-puberulent in inflorescence. Stem dark red, very thick, middle 
part 4-15 mm thick, base usually clavate; covered with numerous, 
dense, ovate or ovate-lanceolate, up to 15 mm long, scales. 
Inflorescence cylindrical or, rarely oval, fairly dense, usually longer 
than the highly shortened remaining part of stem, with thick glabrous 
axis. Bract scales ovate to broadly lanceolate, sometimes highly 
enlarged and flattened, glabrous or very sparsely puberulent. Calyx 
segments free, usually incised up to almost middle into 2 lanceolate 
teeth, up to 15 mm long, brownish, with darker but faint, veins. Corolla 
16-22 mm long, tubular-campanulate, usually brownish or chestnut 
colored when dry, glabrous or subglabrous (with extremely short hairs 
in limb) outside, margin more or less short—fimbriate, its longitudinal 
dorsal line concave (mildly in lower half). Filaments and style 
subglabrous. Anthers glabrous. May to July (Plate VI, fig. 2). 

Parasitic on species of genus Salvia L., predominantly on S. 
glutinosa L. Forests, scrub, rubbly slopes —Caucasus: All regions, 
besides Talysh. General distribution: Armenia-Kurdistan. Described 
from Armenia. Type in Berlin. 

Note. In the southern part of Transcaucasia can be found also the 
closely related species O. anatolica Butss. et Reut. [in Pinard, FI. 
Cariae exsicc. (1843) and in DC. Prodr. XI, 17], differing in having a 
profuse, usually in part whitish lanate inflorescence. Although the name 


114 


108 


O. colorata C. Koch has priority over O. anatolica Boiss. et Reut. 
(diagnosis of this species was published only in 1847), the latter name, 
starting with Boissier, was also accepted by all authors of the flora of 
the Caucasus for O. colorata C. Koch. 

In Flora Rossica (III, 315), another species of the group Arcutae 
G. Beck (O. rapum genistae Thuill.) was reported for the territory of 
USSR, apparently by mistake. The latter grows in the western part of 
Europe (eastward to Austria and Denmark) and is parasitic on species 
of the genera Cytisus L. and Genista L. 


GENUS 1374. Boschniakia! C.A.M. 


C.A.M. ex Bongard in Mém. Akad. Sc. Pétersb. 6 sér. II (1833) 157; 
Reut. in DC. Prodr. XI, 39; Ldb. FI. Ross. III, 323; G. Beck in 
Pflanzenfam. IV, 3b (1895) 132 and in Pflanzenr. IV, 261 (1930) 326. 

Flowers in dense, spicate inflorescence, sessile or subsessile, with, 
or without, 2 bracteoles on sides of calyx, and with single bract scales. 
Calyx gamosepalous, obliquely cupuliform, unequally 2—4 toothed. 
Corolla broadly tubular, inflated, constricted below throat, bilabiate; 
upper lip erect, sinuate or entire, usually several-times longer than 
very short, 3-lobed lower lip. Stamens inserted in corolla tube and 
exserted considerably from its throat; anthers glabrous, anther lobes 
obtuse at both ends. Ovary unilocular, placentae 2—4. Style emerging 
from throat of corolla, bent toward front at apex; stigma bulbous, 
concave in middle or almost bifid. Capsule bivalved. Seeds minute, 
numerous. Perennial herb with thick, simple stems, covered with 
alternate scales. 

Of the two species of this genus, one is found in the USSR, Japan 
and in northwestern parts of North America, the other in China. 


1. B. rossica (Cham. et Schlechtend.) B. Fedtsch. in Fedch. and 
Fler. Fl. (1910) 896, fig. 875 and in Tr. Bot. sada, XXXI, 1 (1912- 
1915) 178; Kom. Fl. Kamch. III, 90; Hulten, Fl. Kamtch. IV, 126; 
Kryl. Fl. Zap. Sib. X, 2547.—B. glabra C.A.M. ex Bongard in Mém. 
Acad. Sc. Pétersb. 6 sér. II (1833) 157; Reut. in DC. Prodr. XI, 39; 
Ldb. FI. Ross. III, 323; Turcz. Fl. baic.-dahur. II, 2, 385; A. Gray, 
Synopt. Fl. North. Amer. II, 1, 313; G. Beck in Pflanzenfam. IV, 3b, 
124, f. 56 (G) and in Pflanzenr. IV, 261, 327, f. 23 (A—G); Kom. FI. 
Manchzh. III, 470; Kom. and Alis. Opred. rast. Dalnevost. kr. II, 936; 
Govorukhin, Fl. Urala, 464, fig. 152.—Orobanche rossica Cham. et 


'Named after A.K. Boschniak, an amateur botanist and horticulturist. 


mt 


~ 


109 


Schlechtend. in Linnaea, III (1828) 32.—O. glabra Hook. FI. bor. 
amer. II (1940) 91 tab. 167.—Ic.: Gmelin, Fl. sib. III, tab. 45, f. 2; 
Hook, I.c.; G. Beck, l.c.; Miyoshi and Makino, Pock.-atlas alp. pl. Jap. 
II, tab. 66, f. 379; Sugawara, Ill. fl. of Saghal, IV, tab. 768; Govorukhin, 
lke 

Perennial. Plant 15—40 cm tall, subglabrous. Stems often in groups 
of 2-8 from clavate base, middle part 6-15 mm thick, glabrous, 
brownish or reddish, covered with numerous, broadly oval or deltoid, 
5-11 mm long, scales. Inflorescence cylindrical or oblong, thick and 
many-flowered, usually one and a half times as long as remaining part 
of stem. Bract scales broadly ovate, obtuse or subacute, equaling or 
somewhat exceeding calyx, glabrous, or ciliate along margin. Bracteoles 
absent. Calyx 2 times as short as corolla, obliquely cupuliform, with 
3-5 unequal, short, teeth, 3-4 toward front longer, toward back one 
small tooth, extremely short or else entirely absent, margin usually 
short lanate. Corolla broadly tubular 10-12 mm long, brownish with 
reddish, distinctly inflated from throat to base, with short limb, along 
margin usually short-lanate; upper lip entire or mildly sinuate, concave, 
usually 3—4 times as long as very short, 3-lobed lower lip. Filaments 
inserted 2-3 mm above base of corolla tube, glabrous or subglabrous 
along with anthers emerging from throat of corolla. Anthers glabrous. 
June to July (Plate VI, fig. 3). 

Parasitic on species of Alnaster Spach, predominantly on A. 
fruticosus (Rupr.) Ldb. Scrub.—Arctic Region: Arctic Europe (eastern 
part), Arctic Siberia, Chukhotka, Anadyr. European USSR: Dvina- 
Pechora (eastern part). Western Siberia: Ob’ Region, Altai. Eastern 
Siberia: All regions. Soviet Far East: All regions except Ussuri. General 
distribution: Japan, Bering Sea Region, North America (northwestern 
part). Described from Kotzebue Bay and Chamisso Island in Bering 
Strait. Type in Leningrad. 


GENUS 1375. Mannagettaea’” H. Smith 


H. Smith in Acta Horti Gotoburg. VIII (1933) 135. 

Flowers in dense, corymbose, almost capitate inflorescences, sessile 
or on extremely short pedicels, in axils of bract scales and with 2 
bracteoles on sides of calyx. Calyx gamosepalous, tubular, 4—5S-toothed. 
Corolla erect, tubular, bilabiate, upper lip entire, galeate; lower lip 3- 
lobed, with 3 almost identical lanceolate-linear lobes. Stamens inserted 


‘Treatment by M.G. Popov. 
In honor of monographer of family Orobanchaceae, G. Beck von Mannagetta. 


116 


110 


in corolla tube. Anthers glabrous or hairy, anther lobes mucronate at 
tip. Ovary unilocular; style with globose stigma. Capsule bivalved. 
Seeds tiny, numerous. Perennial herb with extremely short, usually 
underground stems, covered with few scales. 

Two of 3 species of this genus occur in China; the third has recently 
been described from the Sayan Mountains by M.G. Popov. 


1. M. ircutensis M. Pop. in Bot. mat. Gerb. Bot. inst. AN SSSR, 
XVI (1954) 10, fig. 2.—Ic.: M. Popov. l.c. 

Perennial. Plant 1—7 cm tall, subglabrous, violet-brown, densely 
pulvinately many-stemmed. Stems highly shortened, usually hidden 
underground, neither thickened nor branched at base, with numerous 
ovate, alternate or almost opposite scales. Inflorescence corymbose, 
almost capitate, with 4-15 dark violet flowers, clustered together, 
sessile or on extremely short pedicels. Bract scales broadly ovate or 
oblong, obtuse, margin ciliate, with slightly curly hairs. Bracteoles 
linear, somewhat shorter than calyx. Calyx about 20 mm long, tubular, 
5-toothed, teeth 4-5 times as short as calyx tube, ovate, subacute, 
margin densely ciliate with short curly hairs. Corolla about 25 mm 
long, tubular, somewhat inflated in lower half, limb slightly patent, 
bilabiate, upper lip galeate, entire, obtuse, margin densely ciliate with 
curly hairs, lower lip incised to base into 3 lanceolate-linear lobes, 
ciliate with curly hairs. Filaments glabrous or subglabrous; anthers 
concealed under galea. Stigma globose. July to August (Plate VI, fig. 
4). 

Parasitic on Caragana jubata (Pall.) Poir. In open spruce forests.— 
Eastern Siberia: Angara-Sayan (Eastern Sayan). Endemic (?). Described 
from Buryat-Mongolian ASSR (Tunkin aimak, near village of Turan, 
in riparian spruce forest, 22 VII 1953, L.V. Bardunov). Type in 
Leningrad. 


GENUS 1376. Phacellanthus! Sieb. et Zucc. 


Sieb. et Zucc. in Abh. Akad. Miichen, IV, 3 (1846) 141; Reut. in DC. 
Prodr. XI, 720; G. Beck in Pflanzenfam. IV, 3b (1895) 132 and in 
Pflanzenr. IV, 261 (1930) 332. 

Flowers in corymbose, almost capitate inflorescences, on short, 
thick pedicels, in axils of bract scales, without bracteoles. Calyx with 
2 lateral, caducous, linear calyx lobes. Corolla tubular; slightly 
constricted at place of insertion of stamens, with long tube, smoothly 


‘From the Greek phacelos—cluster, and anthos—flowers. 


117 


111 


bent in front, bilabiate; upper lip bilobed, with short broad lobes, 
lower lip 3-lobed, with short, ovate lobes. Stamens concealed in corolla 
tube; anthers with broad connective and anther lobes obtuse at both 
ends. Ovary unilocular, with 3 carpels, 6 placentae; style clavate at 
tip; stigma slanting, somewhat bilobed. Perennial herb with short, 
often underground stems, emerging in large numbers from thin, 
branched rootstock and covered with ovate obtuse scales. 
Monotypic genus. 


1. P. tubiflorus Sieb. et Zucc. in Abh. Akad. Miinchen, IV, 3 
(1846) 141; Reut. in DC. Prodr. XI, 720; Matsumura, Ind. pl. japon. 
II (1912) 576; G. Beck in Pflanzenr. IV, 261, 332, f. 24 (A-E).—P. 
continentalis Kom. in Izv. AN SSSR, VII ser. No. 3 (1930) 273, fig. 
1, 2 and 3; Kom. and Alis. Opred. rast. Dalnevost. kr. II, 935; 
Sugawara, Ill. Fl. of Sachal. IV, tab. 769.—Ic.: G. Beck. l.c.; Kom., 
l.c.; Sugawara, l.c. 

Perennial. Plant up to 18 cm tall, glabrous. Stems short, often 
underground, not thickened at base; scales fairly numerous, obtuse, 
ovate or ovate-lanceolate, upper ones longer, usually 10-15 mm long. 
Inflorescence, corymbose, almost capitate, usually with 2-10 flowers. 
Flowers on extremely short, thick pedicels, colored with citrous aroma. 
Bract scales ovate or lanceolate, obtuse or acute at apex, usually shorter 
than corolla tube. Calyx of 2 (very rarely 3) free, narrowly linear, 
caducous calyx lobes, resembling bracteoles. Corolla tubular, white, 
22-35 mm long, bilabiate. July to August (Plate VI, fig. 5). 

Host plant not precisely determined; reported on Cimicifuga 
dahurica Maxim. and Deutzia parviflora Bge. 

Coniferous and mixed forests.— Soviet Far East: Ussuri (southern 
part), Sakhalin (southern part). General distribution: Japan, China. 
Described from Japan. Type in Miinich (7). 

Note. The differences of the continental plants compared to the 
Japanese ones (smaller flowers and longer, obtuse bract scales) cited 
by V.L. Komarov (1930) do not hold up and, in our opinion, are not 
consequential. 


Family CXLVII. LENTIBULARIACEAE’ 
LINDL. 


Flowers zygomorphic, monoecious. Calyx bi- or 5-fid, persistent 
in fruit. Corolla bilabiate, with short tube spurred at base; upper lip 


‘Treatment by E.I. Steinberg. 


118 


J 


bilobed, lower 3-lobed, or lobes indistinct. Stamens 2, with flat, 
spreading filaments, inserted at base of the upper lip of corolla; anthers 
uni- or bilocular. Carpels 2, forming superior unilocular ovary with 
numerous ovules on free central placenta; style extremely short, with 
bilobed stigma, one lobe considerably longer than the other, flatter, 
the other sometimes not developed. Fruit unilocular many-seeded 
capsule, dehiscing by 2 valves or rupturing irregularly. Seeds minute, 
nonendospermic. 

Perennial insectivorous herbs with leaves entire or cut into filiform 
segments. 


Tribe. UTRICULARIEAE Kamienski in Zap. Novoross. Obshch. 


est. XV, 1 (1890) 210; Pflanzenfam. IV, 3b, 118.—Flowers solitary 


or in racemose inflorescence. Corolla bilabiate, possessing a distinct 
spur. Ovary with many ovules, on central placenta. Capsule with 
persistent, somewhat connate calyx and persistent stigma, orbicular or 
ovate-oblong. Terrestrial, marshy or aquatic herbs. 


KEY TO GENERA 


1. Marshy or terrestrial plants with radical rosette of entire leaves. — 
Flowers solitary at tip of leafless peduncle. Plant with true roots 
ae pee ened 5, eos tone tual op ea hentai 1377. Pinguicula L. 
+ Aquatic or marshy plants with leaves, incised into linear segments, 
bearing hollow bladders. Flowers in racemes. Plant without roots 
sea vivedbvatbvies oll calSante URES steed a aaeas cca Die 1378. Utricularia L. 


GENUS 1377—Pinguicula’ L. 


L. Sp. pl. (1753) 17. 

Calyx 5-fid, almost bilabiate. Corolla with expanded throat, with 
bilobed, shorter upper lip, longer, 3-lobed lower lip, tapered into spur. 
Stamens 2, short, with thick filaments and transversely attached anthers; 
anthers unilocular. Stigma subsessile, bilobed, with unequal lobes of 
which upper one is flat, orbicular, with fimbriate margin projecting 
above anthers, lower one small, sometimes undeveloped. Fruit bivalved 
capsule. Marshy or terrestrial plant with rosette or radical, entire leaves 
and with 1 or several single-flowered scapes, emerging from middle of 


'Diminutive form of the Latin pinguis—greasy, so-called because of shiny, as 
though greasy, leaves. 


119 


113 


rosette. Upper leaves covered with small glands and their margin 
becomes involuted because of sensitivity to tiny insects that appear 
around their margin. 

The genus includes nearly 30 species, differing slightly from one 
another. They are distributed in Europe, northern Asia and North 
America to the northernmost limits. Based on the color and form of 
the corolla, as well as the size of the spur, De Candolle divided the 
genus Pinguicula L. into three sections. The species growing within 
the USSR belong to section Pinophyllum DC. 


Section. PINOPHYLLUM DC. Prodr. VIII (1844) 28.—Corolla blue, 
violet, white, pale yellow, variegated, almost 5-lobed, with distinct 
upper and lower lips. Corolla tube oblong or infundibuliform. Spur 
shorter than remaining part of corolla. 


1. Plant, with 1 rosette of 1-3 peduncles. ........ eee eeceseeesseeeeeeee 2. 
+ Plant, forming small mats of 34 rosettes ... 1. P. variegata Turcz. 
2.  Peduncles (scapes) with simple or glandular-pubescence ........... 3. 
+ Peduncle glabrous. Flowers light yellow ................. 2. P. alpina L. 
3. Peduncles pubescent with simple hairs towards base ................ 

AS ATID, SE, RIDIN, AREER NOE 3. P. villosa L. 
+  Peduncles uniformly glandular-pubescent .................sceseeeseeeeeeee 4. 
Ae HE GTOMa Garkeviolet see A A EM Ss 
re CorollalightviGlets Augen. Ae ne A RR Se 6. 
5. Corolla dark violet, with white lobes .... 4. P. bicolor Woloszczak. 
+ Corolla dark violet. Peduncles at time of flowering covered densely 


by short glandular hairs; after flowering hairs fall and peduncles 
become almost glabrous .............:csscesseeeseeereeeeeees 5. P. vulgaris L. 
6. Corolla light violet. Radical leaves surrounded by yellow (dry) 
scales. Pedicel densely glandular-pubescent ..................:csssssssssssseseeees 
BARAT WER REE 1, SRR 6. P. glandulosa Trautv. and Mey. 
+ Corolla light violet. Radical leaves not surrounded by yellow scales 
aAUDASE Aye RE 7. P. spathulata Ldb. 


1. P. variegata Turcz. in Bull. Soc. Nat. Mosc. XIII (1840) 77; 
Ldb. FI. Ross. III, 4; DC. Prodr. VIII, 32. 

Perennial. Plant forming small mats of 3-7 rosettes, 3-6 cm tall 
in flowering stage, finally reaching 8-9 cm. Leaves green, glabrous, 
orbicular-spatulate, up to 8 mm long, 4-5 mm broad, with small acute- 
angled notch at tip and with cilia at base of lamina. Peduncles (scapes) 
glandular-hairy, especially in upper part. Flowers 6 mm long. Calyx 
lobes ovate-lanceolate, 1 mm long, 0.8 mm broad. Corolla tube 
infundibuliform, upper and lower lips unequal, upper bidentate; middle 


120 


114 


part of lower lip spatulate, obtuse, indistinctly toothed, with prominent 
veins, considerably larger than lateral lobes; spur conical, yellow, 2 
times as short as corolla; middle part of corolla together with the 
upper lip pale blue; teeth of corolla lobes white, lower lip also white, 
with yellow spot in center. Fruit 2.5 mm long capsule. June to July. 

In moist places.— Eastern Siberia: Lena-Kolyma. Soviet Far East: 
Okhotsk. Endemic. Described from Yakutia. Type in Leningrad 
(collected by N. Kuznetsov between Aldan and Amakh-Yun rivers in 
1935). 


2. P. alpina L. Sp. pl. (1753) 17; DC. Prodr. VIII, 31; Ldb. FI. 
Ross. III; Schmalh. Fl. I, 299, Kryl. Fl. Zap. Sib. X, 2553.—P. 
flavescens Flérke apud Hoppe, Taschenb. (1800) 119.—Ic.: Hegi, Ill. 
Fl. VI. 1, tab. 244, f. 3; f. 93; Rchb. Ic. Fl. Germ. XX, tab. 1821; 
Fedch. and Fler. Fl. 897.—Exs: GRF, No. 1371; Fl. Hung. exs. No. 
396; Pl. Pol. exs. No. 155; Fl. exs. austro-hung. No 3693; Schultz, 
Herb. norm. No. 937; Pl. Finl. exs. No. 356a; Fl. italica exs. No. 654. 

Perennial. Rootstock extremely short, with extremely, short root 
hairs. Leaves sessile, oblong-obovate, obtuse, entire, involute along 
margin, glandular-viscid, yellowish-green, 1.5—3 cm long, 6-12 mm 
broad. Peduncle (scape) 1 or 2, rarely 3, wholly glabrous, 5-12 cm 
long. Inflorescence erect, rarely nutant. Calyx bilabiate, 3 mm long; 
calyx lobes deltoid; acute. Corolla spurred, yellow, 12-15 mm long, 
white or light yellow, with yellowish spots at base of lower lip, spur 
much shorter than corolla, orange-yellow or greenish, conical, bent in 
front. Capsule 7-9 mm long, acuminate, erect, exceeding calyx 2 times 
in length. Seeds brownish, 0.75—1 mm long. June to August. 

In arctic polar region on shingles, rubbly banks of rivers, in moist 
shady places of calcareous deposits, outcrops, in willow-moss 
marshes.—Arctic Region: Karelia-Lapland, Dvina-Pechora, Baltic 
Region. Volga-Kama, Upper Dniester. Western Siberia: Ob’ Region. 
Eastern Siberia: Yenisei, Lena-Kolyma, Angara-Sayan. General 
distribution: Arctic Region, Scandinavia, Central and Atlantic Europe, 
eastern Mediterranean, India-Himalayas, Bering Strait, North America. 
Described from Lapland. Type in London. 


3. P. villosa L. Sp. pl. (1753) 17; Ldb. FI. Ross. III, 4; Kom. FI. 
Manchzh. II, 474; Kom. Fl. Kamch. III, 93; Kom. and Alis. Opred. 
rast. Dalnevost. kr. II, 941; Kryl. Fl. Zap. Sib. X, 2553.—Iec.: Taliev 
in Tr. Obshch. ispyt. prir. Khar’k univ. XXX, VIII (1904) 120.— Exs.: 
Pl. Finl. exs. No. 357a. 

Perennial. Small, often concealed in peat moss vegetation, 4—10 
cm tall. Leaves up to 1 cm long and 5-8 mm broad, ovate or broadly 


121 


115 


elliptical, entire, horizontally spreading, with small acute-angled notch 
at apex, covered by small glands; lamina continued into broad, fimbriate 
petiole, 2 times as short as lamina, rarely almost as long. Scapes 3- 
10 cm long, usually solitary, rarely 2-3, with fairly long, soft, distant 
hairs towards base, glandular-hairy in upper part. Inflorescence nutant. 
Calyx about 1.25 mm long, glandular-hairy with deltoid, acuminate 
lobes. Corolla bluish violet with darker veins and yellowish spots at 
base of lower lip, white hairs in throat; flowers 7-10 mm long including 
spur; spur subulate, erect, obtuse. Capsule orbicular-ovoid, about 2.5 
mm long, dehiscing from above by 5 obtuse, almost semiorbicular 
teeth. (June) July to August. 

In moist sphagnum bogs, on moist moss tundras, along wet banks 
of rivers and streams.—Arctic Region: Arctic Europe, Arctic Siberia, 
Chukhotka, Anadyr. European USSR: Karelia-Lapland. Dvina-Pechora. 
Western Siberia: Ob’ Region. Eastern Siberia: All regions. Soviet Far 
East: Zeya-Bureya, Uda River area., Kamchatka. General distribution: 
Arctic Region, Scandinavia, Bering Strait, North America. Described 
from Lapland. Type in London. 


4. P. bicolor Woloszczak in Oesterr. Bot. Zeitschr. XX XVII (1887) 
80.—Ic.: Zurzycki in Fragm. florist. and geobot. I, f. 6b, tabl. I, 11, 
14; tabi. II, 17. 

Perennial. Plant acaulous, with 2 peduncles about 20 cm tall, 
uniformly covered with short stalked glandular hairs. Roots fibrillar. 
Leaves in radical rosette, oblong or oblong-lanceolate, obtuse, tapered 
from below, sessile. Calyx campanulate, bilabiate, lower lip bilobed, 
upper trifid up to middle, lobes obtuse. Corolla bilabiate, abruptly 
narrowed into erect, subulate-conical, 5 mm long spur; upper lip 
somewhat shorter than lower, bifid, lower lip trifid, middle lobe of 
lower lip 2 times as long as lateral, equaling spur in length; all lobes 
of corolla white, remaining part of corolla violet. Fruit a capsule larger 
than that in P. vulgaris. Seeds normally cylindrical. 

In marshy meadows.— European USSR: Upper Dniester. General 
distribution: Central Europe (Poland). Described from environs of 
Podmonasterets (Ad pagum Podmonasterz in agro Leopolitano). 
Endemic. Type in Cracow. 


5. P. vulgaris L. Sp. pl. (1753) 17; Ldb. Fl. Ross. III, 2; Schmalh. 
Fl. II, 298; Hulten. Fl. Kamtch. IV, 127; Grossh. Fl. Kavk. IV, 14; 
Kryl. Fl. Zap. Sib. X, 2552.—P. macroceras Willd. ex Ldb. Fl. Ross. 
III (1847-1849) 3; Chamisso in Linnaea, VI, 568, nom.—P. 
macroceras Chamisso in Linnaea, VI (1831) 568; Ldb. Fl. Ross. III, 
3.—P. vulgaris € macroceras Herder in Tr. SPb. bot. sada, I (1872) 


122 


116 


380; Kudo, Fl. Paramush, 159.—Ic.: Rchb. Ic. Fl. Germ. XX, tab. 
1819; Fl. Dan. I, tab. 93; Fedch. and Fler. Fl. 972; Hegi, Ill. Fl. VI, 
1, tab. 244, p. 156, 158, 160.—Exs.: GRF, No. 426; Herb. Fl. Ingr. 
414; Schultz, Herb. norm. No. 938; Pl. Pol. exs. No. 154; Fl. Hung. 
exs. cent. IV, No. 317; Pl. Finl. exs. No. 355. 

Perennial. Rootstock very short. Leaves oblong-elliptical, obtuse, 
narrowed at base, subsessile, light green on upper surface,-glandular- 
viscid, 2-4 cm long, 0.6—2 cm broad, forming a dense radical rosette. 
Scape solitary or 2—3 together, 5-17 cm long, when young almost 
uniformly covered by extremely short glandular hairs, disappearing 
after flowering, rendering peduncle subglabrous. Flowers nutant. Calyx 
3-4 mm long, with sparse, short glandular-pubescence, calyx lobes 
ovate or oblong-elliptic, mucronate. Corolla blue-violet, 15-20 mm 
long including spur, covered with fairly long whitish hairs in throat; 
spur subulate, usually obtuse, almost 2 times as short as remaining part 
of corolla. Capsule ovoid-globose. Seeds 0.7 mm long, 0.1 mm broad, 
light brown. May to August. 

In boggy, hammocky meadows, spring-fed bogs.—Arctic Region: 
Arctic Europe. European USSR: Karelia-Lapland, Dvina-Pechora, 
Baltic Region, Ladoga-IImen, Volga-Kama, Upper Dnieper, Upper 
Dniester. Caucasus: Ciscaucasia. Western Siberia: Ob’ Region. Eastern 
Siberia: Lena-Kolyma, Angara-Sayan. Soviet Far East: Kamchatka 
(south), Sakhalin (Kuril Islands). General distribution: Arctic Region, 
Scandinavia, Northern Europe, eastern Mediterranean, Balkans-Asia 
Minor, Armenia-Kurdistan, Bering Strait, North America. Described 
from Western Europe. Type in London. 


6. P. glandulosa Trautv. et Mey. Fl. ochot. in Middendorff’s Reise, 
1, 2 (1856) 65.—Ic.: Trautv. and Mey. l.c. tab. 31. 

Perennial. Cluster of radical leaves surrounded by scales; scales 
oblong-elliptical, obtuse, yellow (when dry), glabrous. Leaves spatulate, 
7-12 mm long, upper part almost orbicular, obtuse, glabrous, middle 
part narrowed down to base, lower part fimbriate. Scape solitary, 5— 
10 cm long, covered by sessile glands. Calyx 1.5-2 mm long, covered 
with glands outside; upper lip deeply tridentate, teeth uneven, ovate, 
middle tooth smaller, softly apiculate, lateral teeth obtuse; lower lip 
bifid, lobes elliptical, obtuse. Corolla about 10 mm long including 
spur, pale violet, middle part with darker violet veins, inner surface of 
upper lip and spur yellow; lower lip with unequal lobes, middle lobe 
almost square, constricted at bottom, end obtusely truncate, lateral 
lobes extremely broadly ovate, considerably smaller than middle one; 
lobes of upper lip ovate-oblong, throat bearded; spur erect, conical, 
obtuse, 1-2 mm long. Capsule orbicular-ovoid, 2—2.5 mm in diameter. 
June to August. 


123 


117 


In sphagnum and clayey bogs.—Arctic Region: Anadyr. Eastern 
Siberia: Lena-Kolyma. Soviet Far East: Okhotsk, Uda River.area. 
Endemic. Described from Uda River estuary. Type in Leningrad. 


7. P. spathulata Ldb. in Mém. Acad. Sc. Pétersb. V ( 1815) 575; 
Roem. and Schult. Syst. veg. I, 192; Ldb. Fl. Ross. III, I (1847-1849) 
Bi A 

Perennial. Leaves in radical rosette; 3-4 in number, orbicular- 
spatulate, obtuse, entire; 1.2 cm in diameter, upper surface of lamina 
sparsely hairy, lower part ciliate along margin. Peduncle (scape) 
solitary, 6-10 cm long, glandular-hairy, especially in upper part. 
Flowers 1—1.5 cm long. Calyx lobes broadly lanceolate, obtuse. Corolla 
light violet with dark lines on surface, spur yellow; throat hairy. June. 

Eastern Siberia: Lena-Kolyma (Dzhegedal, collector Stubendorff). 
Soviet Far East: Okhotsk (Gizhiga). Endemic. Described from Eastern 
Siberia. Type in Leningrad. 

Note. The plants collected by Voronin and Lebedeva in 1914 
perhaps belong to this species (Vitim uplands in former Barguzin 
district). 


GENUS 1378. Utricularia! L. 


L. Sp. pl. (1753) 18, Kamienski in Pflanzenfam. IV, 3b (1895) 119. 

Calyx bilabiate, divided to the base. Corolla bilabiate, with 
extremely short tube and spur, upper lip entire or slightly sinuate 
towards apex, lower larger, entire or indistinctly 3-lobed, swollen in 
middle with a gibbous palate within corolla. Stamens 2, falcately 
declinate, filaments flat; anthers oblong, bilocular, with disjunct or 
coalescing locules, inflexed. Ovary superior, unilocular, of 2 carpels, 
with central plancenta and many ovules; style extremely short; stigma 
bilabiate, sometimes lower lip platelike. Capsule unilocular, dehiscing 
irregularly. Seeds minute, nonendospermic, embryo with extremely 
short cotyledons or without them. Flowers in few flowered cyme on 
leafless, erect peduncles, with tiny bracts at base of pedicel. 
Overwintering of plant by means of globose buds. 

Aquatic plant, rootless and bearing a greater or lesser number of 
bladder traps on leaves divided into linear or filiform segments. Each 
bladder has an opening covered by a valve opening inward, as a result 
of which a tiny aquatic animal can freely enter the bladder but cannot 


‘From the Latin utriculus—bladder, because of the hollow bladders in leaves. 


124 


118 


get out and, probably, upon dying, serves as food for the plant. 


Species of the genus (more than 200) are distributed mainly in 


tropical and, only partially, the temperate zones. They are completely 
absent in the Arctic. 


Kaminski [Pflanzenfam. IV, 3b (1895)] divided the genus into two 


groups on the basis of the morphology and development of the 
vegetative organs: land forms with entire leaves are unique to the 
tropical zone; aquatic forms with parted or pinnatisect leaves, usually 
are found in the temperate zone. Only representatives of section 
Lentibularia Gesn., belonging to the group of aquatic forms, are found 
in the USSR. 


hi; 


Trapping bladders on special colorless branches with rudimentary 
colorless leaves. Green leaves without bladders, distichous ...... 
balun’, See herk womcA A yt ey hs eG oy Lie yd 1. U. intermedia Hayne. 
Trapping bladders on green leaves, latter not distichous but on all 
Sidesodaiihs. 2. WausieoseY...ut. batoelios. aseele..eth- ere 2. 
Flowers, about 8 (7-11) mm across, spur extremely short, conical, 
lengthequaling breadth 222:...2ncin kel... Dees See 3. 
Flowers 12-22 mm across. Spur much longer than broad ......... 4. 
Lower lip of corolla, with recurved margin, minutely gibbous, not 
covering throat. te eA Oe 2. U. minor L. 
Lower lip of corolla, completely flat, not gibbous .................0. 
CELEB MAR NM Ny ge he oy i plc A A 3. U. bremii Heer. 
Peripheral segments of leaves sparsely covered by fine bristles. 
Trapping bubbles 1-2 mm long. Plant slender ....................... 
guste. crcl Ue ihe ne epilation es. ietaaiens 4. U. japonica Makino. 
Peripheral segments, just like lobules of leaves, sparsely covered 
by cilia. Trapping bubbles 3—3.5 mm long. Plant sturdier ........ 3: 
Upper lip of corolla 2—3 times as long as gibbous palate of lower 
lip. Stamens closer, but anthers non-adherent. Pedicel highly 
elongated after flowering, recurved at acute angle, but not arcuate 
Sis oil stl ped sige wh 8 es abe tle § Cent lene Deals 5. U. major Schmidel. 
Upper lip shorter or slightly longer than palcate. Stamens with 
adherent anthers. Pedicel barely elongated after flowering, arcuate 
JiR. SO cele Seaton re leo ple need teed Pa 6. U. vulgaris L. 


1. U. intermedia Hayne in Schrad. Journ. fiir Bot. I, 1800 (1801) 


18; Ldb. Fl. alt. I, 17; Ldb. Fl. Ross. III, 2; Schmalh. Fl. II, 299; 
Grossh. Fl. Kavk. IV, 14; Kryl. Fl. Zap. Sib. X, 2550.—Ic.: Hayne, 
l.c. tab. 5; Syreistsch. Fl. Mosk. gub. III, fig. 176; Hegi, Ill. Fl. VI, I, 
f. 167.—Exs.: GRF, No. 881; Fl. Finl. exs. Nos. 1342, 1343. 


125 


119 


Perennial. Plant with floating or decumbent stems, 10—30 cm long. 
Leaves dimorphic, distichous on one set of branches, without bladders, 
reniform in general appearance, 5-12 mm long, ternately segmented, 
furcate lobes with linear lobules on the other set of branched leaves 
underdeveloped, filiform, simple or bi- or trifid, not narrowed at ends, 
bearing 3-5 mm long obliquely ovoid bladders; sometimes lobules at 
apex of leaves obtuse (var. Grafiana Koch) or all lobules on leaves 
gradually long acuminate (var. Kochiana Celak.). Peduncles (scapes) 
10-20 cm long, with 2—6 flowers on 1-1.5 cm long pedicels, having 
tiny bracts at base. Calyx lobes ovate; upper ones acuminate, lower 
sinuate at tip. Corolla about 15 mm across, light yellow, with orange 
bands on upper lip and on palate of lower lip, upper lip broadly ovate, 
2 times as long as palate of lower lip; lower lip orbicular, margin 
entire, spur bent toward it end of same length (6-7 mm). Peduncles 
erect in fruit. June to August. 

In grassy and loamy marshes and in shallow lakes.—Arctic Region: 
Anadyr, Chukhotka. European USSR: Karelia-Laplant. Dvina-Pechora, 
Ladoga-IImen, Baltic Region, Upper Volga, Volga-Kama, Upper 
Dnieper, Middle Dnieper, Upper Dniester, Volga-Don, Volga Region, 
Lower Volga, Urals (southern). Caucasus: Southern Transcaucasia. 
Western Siberia: Ob’ Region, Upper Tobol, Irtysh, Altai. Eastern 
Siberia: Lena-Kolyma, Angara-Sayan, Dauriya. Soviet Far East: 
Kamchatka, Zeya-Bureya, Uda River area, Ussuri, Sakhalin (Kuril 
Islands). Soviet Central Asia: Tien Shan. General distribution: 
Scandinavia, Central and Atlantic Europe, western Mediterranean 
(northern Italy), northeastern China, Korea, Japan, North America. 
Described from environs of Berlin. Type in Berlin. 


2. U. minor L. Sp. pl. I (1753) 18; Ldb. Fl. alt. 18; Ldb. Fl. Ross. 
III, 2; Schmalh. Fl. II, 300; Grossh. Fl. Kavk. IV, 15; Kryl. Fl. Zap. 
Sib. X, 2551.—Ic.: Fedch. and Fler. Fl. 897; Syreistsch. Fl. Mosk. 
gub. III, 177.—Exs.: GRF, No. 880. Fl. sauc. exs. No. 941; Fl. exs. 
austro-hung. No. 3095. 

Perennial. Plant decumbent or floating, 5-20 cm long, Leaves 5-— 
12 mm long and broad, tripartite, with repeatedly furcate lobes, lobules 
filiform, glabrous, bearing at places small (1—1.5 mm), obliquely ovoid 
trapping bladders (f. platyloba Meister—leaves with flat, compressed, 
linear lobes, 0.3-1 mm broad). Peduncles erect, 5-15 cm long, with 
2—4 scaly leaves and with 2-7 tiny flowers in a cyme. Pedicels 0.5— 
1 mm long, with tiny, membranous, broadly ovate bracts at base. 
Calyx lobes orbicular, short acuminate or lower lobes obtuse. Corolla 


7-11 mm, light yellow, usually with reddish brown bands on low 


palates, not concealed by throat; upper lip with a notch at apex, equaling 


126 


120 


palate, lower lip ovate, its lateral margin perishing after pollination; 
spur extremely short, conical. Pedicels recurved in fruit. Capsule nutant. 
June to first half of August. 

In grassy marshes, small lakes and canals, standing water.— 
European USSR: Karelia-Lapland, Dvina-Pechora, Ladoga-Ilmen, 
Upper Volga, Volga-Kama, Upper Dniester, Middle Dniester, Volga- 
Don, Upper Dnieper, Middle Dnieper, Volga Region. Caucasus: 
Ciscaucasia, western and eastern Transcaucasia. Western Siberia: Ob’ 
Region, Upper Tobol, Irtysh. Eastern Siberia: Lena-Kolyma. Soviet 
Far East: Kamchatka, Zeya-Bureya, Ussuri, Sakhalin. Soviet Central 
Asia: Kyzyl-Kum (delta of Amu Darya, Lake Greater Uzun-Adyr), 
Tien Shan. General distribution: Scandinavia, Central and Atlantic 
Europe, India-Himalayas, Japan, North America. Described from 
Western Europe. Type in London. 


3. U. bremii’ Heer ex Kollinker, Verz. Phan. Gewichs. Ziir. (1839) 
142; Vizn. roslin URSR, 401.—U. pulchella C.B. Lehm. in Flora (1843) 
786.—Ic.: Rchb. Ic. Fl. Germ. XX, tab. 1825; Hegi, Ill. Fl. VI, 1, 
167.—Exs.: Fl. exs. austro-hung. No. 3283; Fl. exs. de C. Billot, No. 
3883. 

Perennial. Plant floating, shoots differentiated into green aquatic 
and colorless terrestrial forms, the latter fixed in soil, often possessing 
excurrent forms between aquatic and terrestrial types, aquatic shoots 
6-60 cm long, the leaves orbicular or semi-orbicular, 2-20 mm long 
and 3.5—30 mm broad, dichotomously partite with 9-25 linear lobes, 
each leaf with 8-10 trapping bladders; terrestrial shoots, 2.5—6 cm 
long, unbranched or with 1 or 2 lateral branches (because of being 
colorless, herbaria often leave out terrestrial shoots from their 
collections). Peduncle 3-42 cm long, forming lax receme of 2-13 
flowers, 2—5 scaly leaflets in lower part of peduncle. Calyx bilabiate. 
Corolla resembles that of U. major Schmidel, but smaller in size; 
palate rust colored with bands; upper lip entire or retuse, equaling 
palate in length; lower lip orbicular, flat; spur turbinate, short. Fruit— 
small capsules, globose, comparatively few. Winter buds globose or 
slightly ovoid, with smooth surface, 0.5-5 mm thick, bud leaves of 
mature axis leathery, dark, 1.2-7 mm long, and 1.5-12 mm broad, 
weakly dichotomously partite in 2-5 broadly linear, acuminate 
segments. July to September. 

In grassy marshes and canals— European USSR: Upper Dniester. 
General distribution: Central and Atlantic Europe. Described from 
Switzerland. Type in Ziirich. 


‘Named in honor of Swedish entomologist and botanist Oswald Brem (1791- 
1857), who taught in High School at Ziirich. 


127 


121 


4. U. japonica Makino in Tokyo Bot. Mag. XXVIII (1914) 28.— 
M. vulgaris auct. non L. Franch. and Sav. Enum. pl. jap. I (1875) 298; 
Matsumura, Ind. pl. Jap. II, 2 (1904) 579.—Ic.: Makino. l.c. 29, 
Sugawara, Ill. fl. of Sagh. 4 (1940) tab. 773. 

Perennial. Stem floating, usually lacking roots or rarely with few 
unbranched rhizoids, sparsely branched, up to 2 m long and 2.5 mm 
in diameter in the broadest part, but usually shorter and thinner. Leaves 
numerous, ovate, ovate-oblong, or oblong twice or thrice pinnatipartite, 
up to 6.5 cm broad; leaves densely covered with small, short stalked 
bladders in lower part of leaf segments, bladders 1-2 mm, sometimes 
up to 4.5 mm in diameter, marginal segments of leaf subulate-filiform, 
with few bristles. Peduncle erect, 9-20 cm or longer, including 
inflorescence. Scaly leaves on peduncle numerous, compressed, 3-4 
mm long, ovate, acute, entire, membranous, with few veins. 
Inflorescence erect, 2-13 flowered, each pedical at base having bracts 
2-6 times as long as calyx, pedicel not recurved after flowering. Calyx 
about 4 cm long, persistent; upper calyx lobes oval, shortly protruding, 
acuminate, lower lobes slightly shorter, ovate-elliptic, bidentate. Corolla 
bright yellow, up to 2 cm across; upper lip of corolla erect, much 
longer than palate, broadly ovate, lower lip with revolute margin, 
broader than upper lip; palate semicircular, about 7 mm in diameter, 
4 mm high, truncated on top, or depressed, yellow with rust-colored 
irregular lines; spur shorter or almost equaling lower lip of corolla, 
conical, acute or obtuse, slightly recurved at tip. Stamens 2 mm long, 
filaments thick, falcate; anthers orbicular. Pistil shorter than calyx; 
ovary globose, smooth, 1.5 mm in diameter; style thick, erect; stigma 
expanded, orbicular, 1 mm long and broad; ovules numerous, minute. 
Fruit—unknown. June to September. 

In lakes.— Soviet Far East: Sakhalin. General distribution: Japan. 
Endemic. Described from Japan. Type in Tokyo? 

Note. In Makino’s description, the diameter of the bladder is given 
as up to 4.5 mm; however, our study of herbarium specimens of this 
species from Sakhalin and from Japan show that the bladder reaches 
1-2 mm, so that this species sharply differs from U. vulgaris L. 


5. U. major Schmidel in Ic. Pl. ed. 2, I (1793) 80; Syreitsch. FI. 
Mosk. gub. III, 175.—U. neglecta Lehm. Ind. schol. Hamb. (1828) 
38; Schmalh. FI. II, 299; Hegi, Ill. Fl. VI, 1, 167; Vizn. roslin URSR, 
401.—Ic.: Schmidel, l.c. tab. XXI, Syreitsch. l.c. 176. 

Perennial. Plant floating, shoots up to 150 cm long, with distichous 
leaves but projecting in all directions. Leaves divided into 2 or 3 large 
lobes, often even in 4, each lobe repeatedly furcate into numerous 
lobules with many (from 8-25) trapping bladders. Peduncle 6-60 cm 


128 


122 


long, mostly green or brown, with 1-4 scaly leaves. Inflorescence 3— 
12-flowered, lax raceme, pedicel 3—5 times as long as ovate-lanceolate 
bracts, pedicel highly elongated after flowering; often double its length, 
diverging at an acute angle or even horizontally, but not arcuate. Corolla 
mostly larger than in U. vulgaris L., 12-18 mm across, short tubular- 
campanulate, bilabiate, lemon colored, with orange bands of palate of 
lower lip; upper lip 2—3 times as long as palate, ovate or ovate-oblong, 
slightly sinuate at apex, lower lip orbicular, margin nearly flat, not 
recurved. Stamens free, not adhering by anthers. June to August. 

In canals.— European USSR: Upper Volga, Upper Dnieper. General 
distribution: Scandinavia, Central Europe. Described from Germany. 
Type in Miinich. 


6. U. vulgaris L. Sp. pl. (1753) 18; Ldb. Fl. Ross. III, 1; Schmalh. 
Fl. II, 299; Grossh. Fl. Kavk. IV, 14; Kryl. Fl. Zap. Sib. X, 2549.— 
Ic.: Fedch. and Fler. Fl. 897; Syreitsch. Fl. Mosk. gub. III, 175.— 
Exs.: GRF, No. 379; Fl. exs. austro-hung. No. 3282; Herb. Fl. Cauc. 
Nos. 241, 241-b, 241-c; Fl. Hung. No. 1132; Hayek, FI. stir. No. 670. 

Perennial. Plant floating. Stem submerged, 1 cm long or more, 
branched. Leaves 1.5-5 cm long, repeatedly pinnate, divided, with 
thickly linear, fimbriate segments and sufficiently large bladders on all 
leaves, bladders obliquely ovoid, somewhat laterally compressed, 3-4 
mm long. Peduncle erect, 15—30 cm long, having several scaly leaves 
in upper part. Flowers 5-10, medium sized, arranged in raceme. Pedicel 
2-3 times as long as bract. Calyx lobes ovate, upper lobe obtuse, 
lower sinuate. Corolla 12-20 mm across, orange-yellow, with reddish 
brown bands on lower lip; upper lip orbicular-ovate, with recurved 
margin, shorter or somewhat longer than palate; lower lip slightly 3- 
lobed, obtuse, with margin recurved; spur 7-9 mm long, with middle 
part about 2 mm broad, not pressed toward lower lip. Pedicel, reflexed 
after flowering. Stamens joined by anthers. Capsule nutant. Peduncle, 
calyx and spur brownish. June to September. 

In standing and slowly flowing water, in canals and marshes.— 
European USSR: Karelia-Lapland, Dvina-Pechora, Baltic Region, 
Ladoga-IlImen, Upper Volga, Volga-Kama, Upper Dnieper, Middle 
Dnieper, Volga-Don, Trans-Volga, Upper Dniester, Bessarabia, Black 
Sea, Lower Don. Caucasus: Ciscaucasia, western and southern 
Transcaucasia, Dagestan. Western Siberia: All regions. Eastern Siberia: 
All regions. Soviet Far East: Kamchatka, Zeya-Bureya, Uda River 
area. Soviet Central Asia: Aralo-Caspian, Balkhash Region, Kyzyl- 
Kum (Amu-Darya Delta). General distribution: Scandinavia, Central 


129 


123 


and Atlantic Europe, Western Mediterranean, Japan, North America. 
Described from Europe. Type in London. 


Family CXLVII. GLOBULARIACEAE’ LINK 


Flowers bisexual, zygomorphic, in globose head with involucre 
bracts. Calyx persistent. Corolla zygomorphic, bilabiate, upper lip bifid 
and lower longer, 3-partite. Stamens 4, didynamous, epipetalous, lower 
ones longer, staminode absent. Ovary superior, unilocular, with | 
pendulous ovule. Style 1. Fruit single-seeded nut enclosed in calyx. 

This family includes the genera Globularia L. and Cockburnia 
Balf. f., the latter monotypic and endemic to Socotra Island. 


GENUS 1379. Globularia? L. 


L. Sp. pl. (1753) 95; Schwarz. in Bot. Jahrb. 69 (1939) 318. 

Calyx deeply 5-partite, teeth lanceolate, subequal, hairy, similar 
to involucre bracts. Corolla longer than calyx, tubular, bilabiate, upper 
and lower lips split into linear lobes. Stamens and style far exserted 
from corolla tube. Nut single, included in persistent calyx. Flowers in 
conical, globose head, with involucre bracts. Perennial herbs with 
underground, woody caudex, with petiolate radical leaves forming 
rosette and sessile lanceolate cauline leaves. 

This genus includes 22 species, distributed primarily in 
Mediterranean countries. The general range of this genus extends from 
Cape Verde islands, Madeira and Canary in the West to southern 
Arabia, Transcaucasia and Trans-Volga in the east; moreover in the 
South it covers North Africa (from Morocco to Somalia) and in the 
North, the mountains of Central Europe. One species of this genus is 
found in the Baltic Sea islands of Eland and Gotland. 

For flora of the Soviet Union, besides the species described below, 
in the literature there is an old reference by S.P. Krascheninnikov 
(Krascheninnikov and Gorter, Flora ingrica (1761) 20) to a locality of 
G. vulgaris L.: “in dry meadows and fields along Luga River.” Neither 
herbarium specimens nor later records confirm the reference of 
Krascheninnikov . The closest locality of G. vulgaris L. is the Swedish 
island named above (Eland Island), the classical locality of the 
nomenclatural type of this species. 


'Treatment by E.G. Bobrov. 
From the Latin globus—a sphere, because of the shape of the inflorescence. 


130 


124 


1. Caudex without stolons. Leaves entire or emarginate at apex with 
mucro or minutely 3-dentate, middle tooth much smaller. Upper 
cauline leaves lanceolate, acuminate though not attenuated 
(southeastern European part of USSR and Stavropol plateau) ..... 
FEL aS See A ae ORE Re LER AY Seo aie hon: 1. G. aphyllanthes Cr. 

+  Caudex with procumbent stolons. Leaves cuneate-obovate on short 
petioles, obtusely 3-dentate at apex. Upper cauline leaves lanceolate, 
attenuated-acuminate, almost aristate (Great Caucasian Range, 
Transcaucasia, Crimea) .......... 2. G. trichosantha Fisch. and Mey. 


1. G. aphyllanthes Cr. Inst. rei Herb. I (1766) 350; Schwarz in 
Bot. Jahrb. 69, 334.—G. vulgaris auct. fl. Eur. med. ac Ross. nec 
typus Linnaei; Lam. Encycl. II (1788) 730; Schmalh. FI. II, 300, pro 
minima parte—quoad pl. volg. and transvolg; Grossh. Fl. Kavk. IV, 
15, only in Stavropol district—G. vulgaris ssp. willkommii Wettst. in 
Bull. Herb. Boiss. III (1895) 286.—G. willkommii Nym. Sylloge (1854) 
140; Korsh. Tent. Fl. Ross. or. 329; Fedch. in Fl. Yugo-Vost. VI, 
247.—Ic.: Fl. Yugo-Vost, VI, fig. 646; Hegi, Ill. Fl. VI, 1, tab. 245, 
4, Schwarz. l.c. tab. XV, 1-5, XVIII, 26-29, XX, 16. 

Perennial. Plant 5-25 cm tall, Rootstock, short branched. Stems 
ascending, simple, with one head, foliated. Cauline leaves few, distant; 
radical leaves in distinct rosette, spatulate or broadly ovate, gradually 
narrowed into petiole, entire, obtuse or emarginate, with mucro in 
notch; cauline ovate to lanceolate, with narrowed base, sessile, entire, 
acuminate. Heads solitary, terminal, 1.3—1.7 cm in diameter. Involucre 
bracts lanceolate, acuminate, margin fimbriate. Calyx hairy, with 
subulate pointed serration. Corolla grayish blue, 6-8 mm long; upper 
lip deeply bifid with narrowly linear lobes; lower lip somewhat longer 
than upper, trifid with linear lobes. Nut about 1.5 mm long, oblong. 
Flowering June. Fruiting July. . 

On calcareous and chalky slopes, rubbly peaks of hills, in open 
steppe communities and sometimes along edges of scrub. Extremely 
rare plant.— European USSR: Volga-Don (extreme southeast, environs 
of Kamenka in Ulyanovsk district, Lavinsk steppe in Syzran district, 
Zhiguli, environs of Kamenaya Chasha), Lower Don (extreme northeast, 
environs of Khvalynsk, near Nulatka, near Ulyanovka), Trans-Volga 
(Sergievsk, Lipovka, Chershila, vicinity of Bugulma, village of 
Taldybulak, Buguruslan district, Belebeevsk Canton, Kozheevo; 
Sokolov farm and in the town of Ichku of Uralsk district). Caucasus: 
Ciscaucasia (Stavropol plateau, environs of Stavropol). General 
distribution: Central Europe, western Mediterranean—from Asturia 
to Illyria, Balkans (as far as Dobrudzha in the east). Type of species 
comes from the mountains of Italy and Illyria and is illustrated in the 
works of Camerarius in 1588. 


131 


125 


Note. A very rare plant, undoubtedly a relict, its geographical 
range, in our flora, is divided into two parts: one—the aforementioned 
locations in the southeastern European territory, and the other, a small 
part, the vicinity of Stavropol in the northern Caucasus. The main 
geographical range of the species extends from Asturia to Dobrudzha. 
Our localities are separated by a distance of over one-and-a-half 
thousand kilometers from the main European range of the species. 


2. G. trichosantha Fisch. and Mey. in Ind. V Horti Petrop. pro 
1838 (1839) 36; Grossh. Fl. Kavk. IV, 15; Schwarz in Bot. Jahrb. 69, 
333.—G. vulgaris auct. non L.: M.B. Fl. taur.-cauc. I, 91; Ldb. FI. 
Ross. III, 455; Schmalh. FI. II, 300, p.p.; Grossh. Fl. Kavk. IV, 15, 
rast. Zakavk.—G. cordifolia auct. non L.: M.B. FI. taur.-cauc. I, 91; 
Ldb. FI. Ross. III, 455.—Ic.: Jaub. and Sp. Ill. pl. or. II, 259; Schwarz. 
lc. tab. XV, 14-17, XVIII, 22-25.—Exs.: Callier, Iter taur. tert. a. 
1900, No. 710. 

Perennial. Stem 5-30 cm tall. Rootstock multi-headed caudex 
developing woody, procumbent stolons, taking root and becoming the 
starting point of new plants. Plant glaucescent, glabrous. Radical leaves 
in rosette, cuneate-obovate, on short petioles, apex obtuse, 3-dentate 
at tip, middle tooth smaller than lateral ones; cauline leaves sessile, 
from broad to linear-lanceolate, acuminate to attenuated-cuspidate. 
Head solitary, about 1.5 cm in diameter. Involucre bracts lanceolate 
or narrowly lanceolate, attenuated and subulately acuminate, fimbriate. 
Calyx lobes subulately acuminate, 3-4 times as long as tube. Corolla 
lobes extremely long, subfiliform, flexucous, twice as long as tube, 
upper lip almost markedly bifid. Nut about 1.5 mm long, oblong. 
Flowering June. Fruiting July. 

On mountain-steppe rubbly slopes, in Transcaucasian mountains 
up to 2000 m.—European USSR: Crimea (Neizats in vicinity of 
Simferopol, Mangup-Kale, Sudak, Kara-subazar). Caucasus: 
Ciscaucasia (Mt. Oshten in state reserve), eastern Transcaucasia 
(Ermani in south Ossetia, vicinity of Mtskhet), southern Transcaucasia 
(Alagez, Lake Sevan Basin, Daralagez, vicinity of Shami in Suralin 
district, Nakhichevan ASSR; Karakush, Karagut in Nakhichevan district, 
vicinity of Akhur in Norashen district). General distribution: Balkans- 
Asia Minor (from Adrianopol to Kagyzman in former Kars Region), 
Armenia-Kurdistan (?), Iran (Tabriz Region). Described from Anatolia. 
Type in Leningrad. 


132 


126 


Family CXLIX. PHRYMACEAE’ SCHAUER 


Flowers bisexual: Calyx tubular, bilabiate, with 2 broadly triangular 
lower teeth and 3 upper subulate-hooked teeth, with 5 main veins. 
Corolla tubular, with bilabiate limb; upper lip shorter slightly bilobed; 
lower lip longer and broader with 3 ovate lobes. Stamens 4, included 
in tube, with lower pair inserted higher up and longer than upper; 
anthers bilocular. Ovary free, unilocular with 1 erect ovule at base; 
style simple, with bifid stigma. Fruit with a nonendospermic seed, 
enclosed by calyx somewhat enlarged and bent below; cotyledons 
convolute or complex. Flowers in pairs on lax and thin raceme. Bracts 
4 in each flower. Leaves opposite, petiolate. Stem 4-angled. Perennial 
herbs. 

Note. The monotypic lopseed family was segregated from the 
vervain family by Schauer (Schauer in DC. Prodr. XI (1847) 520) on 
the basis of the presence in the genus Phryma of a unilocular ovary, 
with a solitary ovule. Bentham and Hooker included this genus in a 
separate tribe of the vervain family. 

In Engler’s system, the lopseed family is placed between the 
families Myoporaceae and Plantaginaceae. 

There are two works available on the morphology and anatomy of 
the Phrymaceae; the first by Briquet [in Mém. Soc. Phys. and Hist. 
Nat. Gen. (1896) 1], and the second by Holm [in the Bot. Gazetta Vol. 
56 (1913) 306]. These works, however, do not resolve the problem of 
the proper placement of the lopseed family in the phylogenetic system. 

The pollen morphology of the Phrymaceae provides very little 
information but, to some extent, gives a reason for assuming the closest 
relatives of this family to be the Verbenaceae. 

The vervains and lopseeds are similar in the structure of the exine 
of the pollen grains and especially in the presence of the third layer, 
situated between the ektexine and endexine. 


GENUS 1380. Phryma? L. 


L. Sp. Pl. (1753) 601. 

Flowers in pairs on slender, long racemes. Bracts linear-lanceolate, 
4 in each flower. Calyx bilabiate, upper 2 teeth, short, deltoid, lower 
3 long, subulate, hooked. Corolla tubular, bilabiate (5)7—8 mm long, 


‘Treatment by L.A. Kuprianov. 
"Local Indian name. 


133 


127 


upper lip considerably shorter than lower, bilobed, lower broad, 3- 
lobed. Fruit—achene with persistent calyx. 
Plant of broad-leaved forests of East Asia and North America. 


1. P. leptostachya L. Sp. Pl. (1753) 601; Kom. and Alis. Opred. 
rast. Dalnevost. kr. II, 941; Britt. and Brown, Ill. Fl. North Amer. III 
(1913) 244 and Ic. 

Perennial. Rootstock very poorly developed, sometimes entirely 
absent (Holm calls it pseudorhizome). Main root perishes soon, and 
lateral roots form a cluster unbranched roots. Stem erect, 4-sided, 
pubescent, with sparse, long hairs bent below, sometimes subglabrous. 
Leaves long-petiolate, opposite, above and beneath covered by sparse, 
more or less, long hairs and short sessile glands; coarsely serrate- 
dentate along margin, serration simple or doubled, broadly deltoid, 
acute, erect, especially in lower leaves; lower leaves distant, with 4— 
6 cm long petioles, lamina broadly ovate, 6-10 cm long and 5-9 cm 
broad, with regular and cuneate base; middle leaves closer, 8-9 cm 
long and 4 cm broad, at base mildly cuneate or orbicular; upper yet 
smaller. Inflorescence apical, solitary, sparse raceme or in several 
racemes. Bracts linear-lanceolate, shorter than calyx. Flower paired, 
6-8 mm long. Calyx 5 mm long. Corolla white with blue veins, S—8 
mm long. Achene 4—5 mm long and 1.5 mm broad, 4-sided, acuminate. 
Flowering July. Fruiting August. 

Broad-leaved forests.—Soviet Far East: Ussuri. General 
distribution: Manchuria, northern Japan, North America. Described 
from North America. Type in London. 

Note. When describing P. leptostachya, Linnaeus had at hand 
evidently, only the North American material. It was subsequently 
confirmed that this species was also found in Asia in the Himalayas, 
China, Manchuria, Japan and in Far Eastern part of USSR. The 
Himalayan, Chinese and southern Japanese plants differ from the 
Manchurian and Far Eastern ones in having narrower leaves with 
smaller, forward-directed teeth. These plants were described from Japan 
as P. oblongifolia Koidz. [Bot. Mag. Tokyo, XLIII (1920) 400]. Yet 
another species was described from China, P. esquirolii Lev. [Fedde, 
Repert, XII (1913) 534], with large entire leaves and a red corolla. 

Several authors (Rafinesque, Briquet and Holm) have indicated 
that the American plants are not homogenous and can be recognized 
as several species. From the comparatively limited material at our 
disposal, the American Phryma is closest to the plants from northern 
Asia, sO we maintain the name P. leptostachya L. for our Far Eastern 
plants. 


128 


Order 37. PLANTAGINALES ENGL. 


Flowers 4-partite, tubular, regular, bisexual or unisexual. Leaves 
usually spiral. 


Family CL. PLANTAGINACEAE!’ LINDL. 


Flowers bisexual, rarely unisexual, with bracts, generally forming 
spicate, axillary inflorescence. Calyx lobes 4, persistent in fruit, keeled; 
2 anterior calyx lobes often somewhat different from posterior. Corolla 
actinomorphic, tubular with 4-lobed limb, membranous. Stamens 4 
(rarely 1-2 stamens rudimentary), alternating with corolla lobes; 
filaments of stamens long, usually far exserted from corolla. Pistil 
one, consisting of 2 carpels; ovary superior, usually bilocular, with 
one or several ovules in each locule, rarely single ovule; style one. 
Fruit, a capsule with few or many seeds, rarely single-seeded 
indehiscent nut. Annual or perennial herbs, rarely shrubs. Leaves 
without stipules, simple, alternate and usually in compact rosette, rarely 
opposite. 

Only three genera are included in the family: Plantago L., Littorella 
Bergius and Bougneria Decne. 


KEY TO GENERA 


1. Flowers bisexual, in spikes (sometimes short, capitate). Fruit— 
capsule with transverse annular suture, along which upper part of 
capsule separates on ripening of seeds; fruit rarely indehiscent (in 
Ps mdcrocarpa) 2.220. edie, Siok. ee: 1381. Plantago L. 

+ Flower unisexual, monoecious, inflorescence few flowered. 
Staminate flower usually single, pedicellate, with 2(3) pistillate 
flowers at base. Fruit single-seeded, indehiscent nut .............. 
wosdteed LER, Oe meres: Sach aes 1382. Littorella Bergius. 


‘Treatment by Yu. S. Grigoriev. 


134 


135 


129 
GENUS 1381. Plantago! L. 


L. Sp. pl. (1753) 112. 

Flowers with bracts, in spike, sometimes capitate and few flowered. 
Calyx lobes 4, usually free, occasionally, 2 anterior calyx lobes 
somewhat different in shape from posterior lobes. Corolla regular, 
membranous, with cylindrical tube and 4-lobed limb. Stamens 4, 
attached to corolla tube and 4-lobed limb. Stamens 4, attached to 
corolla tube and alternating with its lobes; filaments often long, far 
exserted from corolla tube. Pistil one. Fruit—bilocular (rarely 3- or 4- 
locular because of growth of false septum) capsule with one or several 
seeds in each locule. On maturity of seeds, upper part of capsule 
separating along transverse annular suture and falling off 
(circumscissile) capsule indehiscent only in case of P. macrocarpa, 
there being no annular suture. Annual, biennial or perennial herbs. 
Leaves forming radical rosette, rarely, more or less distant, alternate 
or opposite. 

There are 260 species belonging to this genus, distributed over 
the entire globe, but found-mainly in extra-tropical latitudes. Several 
species are ruderal- weeds, dispersed by human activity over the 
different continents. The best examples are the Eurasian species: P. 
major and P. lanceolata. A species introduced to the USSR, although 
it never really became widespread, is the North American species P. 
aristata Michx., found in the Mukachevo Region of Transcarpathia 
(environs of the village of Shkuratovtsy). 


1. Plant with well-developed, foliate stems. Leaves opposite, bearing 
shortened shoots in axils. (Subgenus 2. Psyllium) ..............0006 Pa 
+ Plant acaulous with leaves in radical rosette (sometimes several 
rosettes), and with leafless scape. (Subgenus 1. Euplantago) ..... 3. 
2. Lower 2 bracts broad at base, orbicular-ovate, suddenly narrowed 
at apex into linear-subulate 3-10 mm long appendage. Upper bracts 
without appendage, broadly obovate or subspatulate, obtuse ..... 
AE IE, ATER BE IS, OREN GE IES OE SOS 26. P. indica L. 
+ Lower bracts ovate or ovate-lanceolate, upper gradually narrowed 
into oblong-linear, sometimes fairly long, appendage. Upper bracts 
similar to lower, rarely broader and with shorter appendage......... 
apawharab es staundebynsassacdecedssuass stew sic tecseMe Oe OMMRMeRO Mod 27. P. psyllium L. 
3. Two anterior calyx lobes connate, forming broad, slightly bilobed 
scale at tip, having 2 herbaceous keels (veins). Posterior calyx 
lobes free. Bracts short or long acuminate ...... 19. P. lanceolata L. 


‘Name formed from planta meaning trace and agere meaning to direct, to lead, 
because leaves of certain species resemble impressions of traces of feet on soil. 


130 


= + 


a+ 


m+ 


Ge) 


136 


Calyx lobes:not:firee coz: ict 0.3.90, SL ee Eo codecs eter ee sede esc 4. 
Leaves broad, ovate or elliptic, with more or less long petioles. 
Spikes narrow, cylindrical, usually rather dense, but interrupted in 
lower part, with more or less scattered flowers; occasionally, flowers 
somewhat scattered over entire length of spike. Bracts shorter 
than calyx, rarely as long or somewhat longer. Capsule with (5)6— 
30(34) seeds. Biennial plant with fibrous root system ................ 
sae, Cire a0, «Bees, «ee Biadina tetas coches pee cats 1. P. major L. 
Different set of:characteristics ..:i.:..niasentla.ton.adas. se 5. 
Capsule 5-7 mm long, without transverse annular suture, 
indehiscent, falling off as a whole. Plant observed in the USSR 
only in Koraginsky Island and Commander Islands in Kamchatka 
REGION: bce.) arse tenets: 5. P. macrocarpa Cham. and Schlechtd. 
Capsule with transverse annular suture; upper na of capsule falling 
Off on» maturity.of, seeds) jeden t! ecanes tele 22 6. 
Corolla tube tomentose or villous or glabrous, in which case corolla 
lobes with long hairs. Leaves narrow, linear or lanceolate sometimes 


pinnatelytlobed: s+. 2icts..:t;-eniernladeatnn te OSS Ge ee Te 
Corolla tube and lobes glabrous ................cssccssssesseeeseeeseseneeeeeeee 10. 
Perennials with well-developed main, often many-headed tap roots. 
Leaves fleshy, canaliculated ...............:sssseseeeees 7. P. maritima L. 
Annvaleplant)..c.: doancbintedacmicads abun eesas ek. bad ee 8. 


Leaves with several scattered, narrow lateral lobes or serration, 

rarely with entire or subentire margin. Posterior calyx lobes rarely 
keeled, winged, winges shortly fimbriate along margin; anterior 
calyx lobes not winged. Capsule with 3-5 seeds ...............::sssseeeee 
l. .Sisnrne eaves d..sepete staiiot. acini tie 6. P. coronopus L. 
Leaves always entire and subentire margin. All 4 calyx lobes without 
wings along keel. Capsule with 2 seeds ...............:scsssceseeseeseeeeeeees 9. 
Corolla lobes glabrous; tube densely villous ................eeeeeeeeeeee 
ait tm hare daterccslacsithen. need be, dees 24. P. lachnantha Bee. 
Corolla lobes villous; tube glabrous ..... 25. P. lagocephala Bee. 
Perennial with well-developed tap root. Leaves lanceolate or linear- 
lanceolate, 0.5—1.5(2) cm broad, with short or fairly long petioles. 
Spikes 2—5(6) cm long, dense, sometimes somewhat interrupted 
only at base. Capsule with 2(4)-5 seeds. Plant of Lena-Kolyma 
Resionas. corms. dite. pas. eager. sake: 16. P. canescens Adams. 
Ditterent set of charactemisGies, |x cee oie cae ant cesses pscbereecesseen Lis 
Perennial 3—15(25) cm tall. Leaves elongated lanceolate, rarely 
elliptic. Young leaves (as well as scapes), densely, mature leaves 
more rarely arachnoid-lanate. Spikes 0.5-4(5) cm long. Capsule 3— 
4 mm long, with 2—3(4) seeds. Plant of high altitude regions of 
Soviet CentralvAStajjcaissasomigis: dateteary sae 11. P. arachnoidea Schrenk. 


137 


12. 


14. 


18. 


19. 


20. 


131 


Different set of characters. Pubescence on leaves usually different, 
orelse:glabrousisets. cok ore. Leth ee 2S A ieee 12: 
Bracts enclosing calyx, broad (their breadth rarely exceeds length), 
variable, at times irregularly shaped, (3.5)4—6(7) mm long. Spikes 
dense, capitate or short-cylindrical. Capsule 4—6(7) mm long, with 
2 seeds. Perennial, high-altitude plant, in USSR—only in Caucasus 


atid injKarélia Aedes 0. woke. 17. P. atrata Hoppe. 
Different, characteristics: ile... eset. eee Ra Ree ee 13. 
Leaves linear or linear-lanceolate, 0.5—8(9) mm broad. Small, 
annual, rarely perennial plant ................cccccssesscesseseesseeseeeeeeeees 14. 


Leaves broader, ovate, oblong-ovate, elliptic or lanceolate, with 
well-developed petioles, rarely subsessile. Perennial plant, rarely 
annualbiennialit s:icces, oreo secs, lnncretly. clare 1 EER. sens 20. 
Perennial high-altitude plant with well-developed tap root. Bract 
herbaceous, membranous only along margin, glabrous. Corolla lobes 
about 1 mm long. Capsule ovoid, 2-3 mm long, 4-locular ......... 
chshastemetonshciemarieter: dement gO e aeee  domegall cased 12. P. komarovii Pavl. 
Annuals, predominantly growing in campestral deserts and 
semideserts, aS well as 1m StepPe .............::cssscesssceesseeesseeeeeesees 15. 
Leaves with several scattered large teeth. Bracts lanate on keel...... 
pbvoteprsescds. todlad. dons Bi aed ner eTAe GA Re 23. P. notata Lagasca. 
Leaves entire or subentire. Bracts glabrous, rarely hairy ........ 16. 
Capsulewith(G)9-36i seedsjccux Leeteeecess i ee eR cae ile? 
Capsulewwith 2/sced se! anccec beak tevin. ad VCRTO BERS, 18. 
Bracts ovate or oblong-ovate, often narrowed at tip or more or less 
long acuminate ................secee 2. P. tenuiflora Waldst. and Kit. 
Bracts broad, orbicular or orbicular-Ovate ..............ccesceeseseeeees 
dropeereatoirls AE. Bessie assorrec sth vers iasnes 3. P. polysperma Kar. and Kir. 
Corolla lobes orbicular-ovate or orbicular, about 2.5 mm long, 


short acuminate Or SUDACULE .............ccesccceesseeeeeeees 20. P. ovata Forsk. 
Corolla lobes oblong-ovate, elliptic or lanceolate, 1-1.5 mm long 
iT 2S abet ee ets ceed ubimmet one mee se. se ke Ok mein. 19. 


Bracts broad, completely enclosing calyx, usually glabrous ........ 
eetca Mel | yo vest eines apse serial eee. ET ARE 22. P. loeflingii L. 
Bracts broad, but not enclosing by calyx, considerably shorter, 
rarely almost as long, glabrous or more or less hairy ................ 
weaned Hemme oul oN spect ston svact stents «Aisidse ating tebe 21. P. minuta Pall. 
Bracts 2—2% times as short as calyx. Spikes 5—20 cm long, fairly 
dense in upper part, more lax and interrupted in lower part. Leaves 
thicker, ovate or ovate-elliptic, with fairly long petioles ............... 
 Uaspeenee sonal a creases haaet Aaasncrenspdinn tg at neea he BRE 4. P. cornuti Gouan. 
Bracts equaling calyx or only somewhat shorter ..................+ 21: 


132 


21. Leaves ovate-lanceolate, ovate or elliptic, gradually narrowed from 
broad base to short and broad or long petiole. Spikes subcapitate 
or cylindrical, 0.7-3(4) cm long. Capsule ovoid, ellipsoid or 
subglobose, about 3 mm long, with 3(4)—7 seeds. Plant found in 
high-altitude meadows of Pamiro-Alal .............::scssceccssesseeeneeeeeeees 
J Jett ieee 18. P. gentianoides Sibth. and Smith. 

+ Different set of characteristics. Plant growing mainly in campestral- 
forest or in steppe region, rarely in mountains ..................0000 22. 

22. Leaves thicker, large, broad, ovate or orbicular-ovate, with 7-11 
veins and long petioles. Spikes 5-20 cm long, with closely-packed 
flowers, compact and thick. Capsule oblong-elliptic, about 3 mm 


long;4-seeded batons ha Nees eee. 15. P. maxima Juss. 
+ Leaves differently shaped, narrower, rarely broadly ovate, but thin 
with short petioles, almost sessile ..............c:ccescesseescessesseeeeeereees 23. 


23. Calyx lobes membranous, anterior 2 with distinct herbaceous keel 
(veins), posterior with barely distinct keel. Leaves 3-veined, rarely 
having another less distinct pair of lateral veins ..............:.:esee00 
tanh... Lewieerer ac. SU SEO 8. P. schwarzenbergiana Schur. 

+ Calyx lobes membranous, but all 4 calyx lobes with distinct 
liesbaceous keels ©... 2052). RIGA EON OAR ales ce 24. 

24. Perennial. Leaves thicker. Younger leaves (just like scape) densely, 
mature leaves sparsely white-hairy. Spikes 2-12 cm long, fairly 
dense, but often with several scattered lower flowers. Capsule ovoid 
or ovoid-cuneate, 3-4 mm long. Plant of Far Eastern coasts........ 
oes ARON BET alas Sea ed be NR sea oe 10. P. camtschatica Link. 

+! o Different set‘of characteristics: ....5...02--csccdazcskelteraveecte ae eta es 25: 

25. Spikes relatively thin, fairly dense in upper part, interrupted in 

lower part, with generally scattered flowers ... 9. P. depressa Willd. 
+ Spikes dense, cylindrical. Flowers somewhat closely-packed all 
alonerleneth Of, Spike i.s...ssiee. ce EE es A Ce .. 26. 

26. Leaves elliptic or ovate (rarely ovate-lanceolate), usually no more 
than 2-2 times as long as broad, on short petioles 3-10 times as 
short as lamina, or nearly sessile. Scape ascending at base, erect 
above. Spike: 2—6(8) cm 1ON ..........cccceecesteesseeeeeees 13. P. media L. 

+ Leaves elliptic (more rarely elliptic-ovate), lanceolate or narrowly 
lanceolate, 22-5 times as long as broad, on petioles almost as long 
as lamina or shorter, though not more than 2-3 times. Scape erect 
or slightly ascending at base ................. 14. P. stepposa Kupr. 


Subgenus 1. Euplantago Harms in Pflanzenfam. IV, 3b (1895) 
370.—Leaves alternate, mostly aggregated in rosette, more rarely 
diffuse. 


139 


133 


Section 1. PoLYNEURON Decne. in DC. Prodr. XIII, I (1852) 694.— 
Ovules 6 or more, seeds minute, often angular. Corolla lobes short, 
more or less reflexed. Rootstock shortened, main root perishing early. 
Leaves broad, with well-developed petioles, more or less distinctly 
demarcated from lamina. Spike narrow and usually fairly long. 


1. P. major L. Sp. pl. (1753) 112; M.B. FI. taur.-cauc. 1, 108; 
Ldb. Fl. Ross. III, 476; Schmalh. Fl. I, 350; O. and B. Fedch. Perch. 
rast. Turkest. V, 193; Grossh. Fl. Kavk. IV, 17; Pilger in Pflanzenr. 
H. 102 (1937) 41; Kryl. Fl. Zap. Sib. X, 2556.—P. sorokini Bge. in 
Tr. SPb. bot. sada, VI (1879) 393.—P. vulgaris Pavlova in Zhurn. 
Russk. bot. obshch. VIII (1923) 95.—P. scopulorum Pavlova, |.c. 96.— 
Ic.: Hegi, Ill. Fl. VI/I, tab. 246 and f. 102; Pilger, l.c. f. 7 and 8; 
Nikitin in Fl. Turkmen. VI, plate XL; Maevsk. FI. (1954) fig. 197.— 
Exs.: GRF, No. 431. 

Biennial. Plant (4)10-50(70) cm tall, with one or often several 
erect or ascending scapes. Leaves broad, ovate or elliptic, obtuse, 
usually rounded at base, entire or somewhat dentate in lower part, 3— 
7 veined, long-petioled, glabrous or sparsely pubescent; petioles 
equaling lamina or longer, rarely shorter. Scapes sparsely pubescent 
with upward directed hairs or glabrous. Inflorescence narrow, 
cylindrical, 2—35(60) cm long, usually fairly dense and interrupted 
only in lower part, with few scattered flowers, rarely flowers scattered 
over entire length of inflorescence. Bracts deltoid-ovate, keeled, with 
broadly membranous margin, usually shorter than calyx, rarely, as 
long or somewhat longer. Calyx lobes 1.5—2.5 mm long, elliptic, obtuse, 
keeled, margin broadly membranous. Corolla lobes ovate or ovate- 
lanceolate, about 1 mm long, usually subobtuse. Capsule ovoid cuneate 
or ovoid (rarely subglobose), considerably (rarely 2 times) longer than 
calyx, with (3)6—30(34) seeds. Seeds about 1 mm long, flattened, often 
slightly angular. (May) June to September (October). 

In ruderal and weedy places, around dwellings, in fields and 
meadows. Widely distributed over the entire territory of the USSR, 
with the exception of the extreme north. More rare in arid zone, and 
occurs there only along river banks and irrigation canals, in abundantly 
irrigated fields, etc. General distribution: Widely distributed in Western 
Europe and Asia; quite common (as adventive plant) in other continents. 
Described from Europe. Type in London. 

Note. 1. Many authors consider that in eastern Asia P. major is 
replaced by a few very closely related species, in particular P. asiatica 
L. and P. japonica Franch. et Savat. Whether these are separate species 
and what characters distinguish them has not at all been resolved. 
V.L. Komarov [Fl. Kamtch. III (1930)] reports for the Kamchatka 


140 


134 


Region only P. asiatica and P. japonica whereas Hultén [Fl. Kamtch. 
IV (1930)] lists only P. major for this region. According to Sugawara 
(Ill. Fl. of Sachal.), both P. asiatica and P. japonica are found in 
Sakhalin, whereas Ohwi [Fl. of Japan, (1953)] reports for Japan P. 
asiatica and P. major var. japonica (Franch. et Savat.) Miyabe. 
According to Pilger [Pflanzenr. H. 102 (1937)], P. asiatica and P. 
japonica have flowers with short pedicels, whereas P. major has sessile 
flowers. Ohwi distinguishes P. asiatica and P. major by other characters 
but treats P. japonica as a variety of P. major. 

Note 2. P. fonticola Kom. [Fl. Kamtch. III (1930) 98], described 
from Kamchatka, evidently, represents only a form of P. major, 
growing along banks of hot springs. I counted up to six seeds in 
capsules of authentic specimens of P. fonticola. According to V.L. 
Komarov there should always be four seeds. 

Economic importance. The leaves of P. major are used in local 
medicine as an antihemorrhagic and for healing wounds. 


Section 2. MicropsyLLiuM Decne. in DC. Prodr. XIII, 1 (1852) 
636.—Ovules 6 or more. Seeds minute. Corolla lobes small. Often 2 
stamens rudimentary. Small annuals with linear or narrowly linear 
leaves. 


2. P. tenuiflora Waldst. et Kit. Descr. and Ic. Pl. rar. Hung. I 
(1802) 37; M.B. FI. taur.-cauc. I, 110; Ldb. Fl. Ross. III, 476; Schmalh. 
Fl. II, 350; O. and B. Fedch. Perech. rast. Turkest. V, 193; Grossh. FI. 
Kavk. IV, 17; Pilger in Pflanzenr. H. 102, 68; Kryl. Fl. Zap. Sib. X, 
2556; Nikitin in Fl. Turkmen. VI, 312.—P. minor Fries, Nov. FI. 
Suec. Mant. II (1839) 12.—P. bungei Steud. Nomencl. ed. 2, II (1841) 
348; Ldb. I.c. 475; Kryl. l.c. 2555.—Ic.: Waldst. and Kit. l.c. tab. 39; 
Pilger, l.c. f. 10; Nikitin, l.c. plate 40.—Exs.: Dorfl. Herb. norm. No. 
5326; Degen. fl. exs. austro-hung. No. 3824; GRF, No. 1130. 

Annual, rarely biennial. Plant 3—20(30) cm tall, with 1 or often 
with several (up'to 15-20) erect or ascending scapes. Leaves linear or 
narrowly linear, 0.5-4 mm broad, subobtuse, broadened at base, 
subentire, rarely remotely dentate margin, somewhat pubescent or 
subglabrous like scape. Spikes cylindrical, narrow, lax (sometimes even 
with few scattered flowers) or short and compact, 0.5—10(15) cm long. 
Bracts ovate or oblong-ovate, somewhat keeled, narrowed toward apex 
or slightly long acuminate, with broadly membranous margin in lower 
part, equaling or exceeding calyx (especially for lower flowers). Calyx 
lobes about 1.5—2 mm long, keeled, broadly membranous, obovate or 
elliptic-ovate, obtuse or short acuminate. Corolla giabrous, with oblong- 
ovate about 1 mm long lobes, corolla lobes sometimes directed upward 


141 


135 


and compactly arranged, concealing style and stamens, having short 
filaments, in other cases corolla lobes more or less reflexed and stamens 
having longer filaments, far exserted from corolla. Capsule much longer 
than calyx, ovate-cuneate, 3—4 mm long, with (6)7—16(20) seeds. Seeds 
fusiform, sometimes curved, slightly angular, 1-2 mm long. May to 
July (August). 

On alkaline soils, takyrs [clay areas in deserts], solonchaks [salt 
marshes], saline meadows, etc.—European USSR: Middle Dnieper, 
Volga-Don, Trans- Volga, Bessarabia, Black Sea Region, Crimea, Lower 
Don, Lower Volga. Caucasus: Ciscaucasia, eastern Transcaucasia, 
Talysh. Western Siberia: Upper Tobol, Irtysh, Altai. Soviet Central 
Asia: Aralo-Caspian, Balkhash Region, mountainous Turkmenia. 
General distribution: Scandinavia (only on Eland Island), Central 
Europe, Balkans-Asia Minor, probably in Dzhungaria-Kashgaria. 
Described from Hungary. Type in Budapest. 

Note. According to N.V. Zinger [Zap. Kievsk. Obshch. est. 19, 1 
(1904)], plants of P. tenuiflora growing under conditions of low 
temperature and high humidity have relatively short scapes, dense 
(capitate) spikes, broad bracts and a smaller number of seeds in the 
capsule. Such plants can survive over winter and again flower and 
bear fruit the following year. On Eland Island, off the western coast 
of Sweden (this locality of the species is isolated from the main area 
of distribution, situated considerably to the south, in steppe and desert 
zones), one finds just such ‘plants. They were described as a distinct 
species.—P. minor Fries, but in reality they undoubtedly belong to P. 
tenuiflora Waldst. et Kit. 


3. P. polysperma Karelin et Kirillow in Bull. Soc. Nat. Mosc. XV 
(1842) 431; Ldb. Fl. Ross. III, 476; O. and B. Fedch. Perech. rast 
Turkest. V, 192; Pilger in Pflanzenr. H. 102, 72; Kryl. Fl. Zap. Sib. 
X, 2555. 

Annual. Plant 2—12(17) cm tall, with one, several or many (up to 
25-30), erect or ascending scapes. Leaves thicker, linear or narrowly 
linear, 0.5—3(4) mm broad, subobtuse, somewhat broadened toward 
base, margin entire. Scapes uniformly sparsely hispid, or glabrous. 
Spikes few-flowered, subcapitate, 2-5 mm long, or many-flowered, 
cylindrical, up to 6—7 cm long, dense, sometimes interrupted only at 
base. Bracts broad, orbicular or orbicular-ovate, somewhat keeled, with 
membranous margin, equaling calyx lobes or somewhat longer. Calyx 
lobes broadly elliptic or ovate, 1.5—2 mm long, obtuse, margin broadly 
membranous. Corolla glabrous, with about 1 mm long ovate lobes. 
Capsule 3-3.5 mm long, ovate-cuneate, much longer than calyx, with 


142 


136 


(7)12-—36 seeds. Seeds fusiform, dark colored, 1-1.5 mm long. May to 
June (July). 

On solonchaks and saline meadows.— Western Siberia: Upper 
Tobol, Irtysh. Soviet Central Asia: Aralo-Caspian (northern part) 
probably north of Balkhash Region. General distribution: Mongolia 
(Gobian Altai), probably in Dzhungaria-Kashgaria. Described from 
Eastern Kazakhstan. Type in Leningrad. 


Section 3. PALEOPSYLLIUM Pilger in Pflanzenr. H. 102 (1937) 75.— 
Ovules 2-4-7. Seeds flattened. Bract and calyx lobes herbaceous, 
broadly membranous along margin. Spike elongated. 


4. P. cornuti Gouan, II]. and Observ. Bot. (1773) 6; Schmalh. Fl. 
II, 350; Pilger in Pflanzenr. H. 102, 83; Kryl. Fl. Zap. Sib. X, 2560.— 
P. gouani Gmel. Syst. Nat. (1791) 251.—P. asiatica auct. non L.; 
Ldb. FI. Ross. III, 1, 479; O. and B. Fedch. Perech. rast. Turkest. V, 
194; Maevsk. Fl. izd. 7, 669; Grossh. Opred. rast. Kavk. 357.—Ic.: 
Pilger, l.c. f. 12.—Exs.: Pilcher, Fl. exs. austro-hung. No. 435. 

Perennial. Plant 20-60 cm tall, with 1-3(5) scapes, usually much 
longer than leaves. Leaves thicker, broad, ovate, or ovate-elliptic, 
subacute or obtuse, gradually narrowed at base into fairly long petiole, 
entire, (5)7-veined, glabrous or hairy along veins beneath; petioles 
also hairy or subglabrous. Scape more or less addressed hairy; spikes 
5-20 cm long, not thick, dense above, lower part lax with scattered 
flowers. Bracts 2—2% times as short as calyx, ovate, or orbicular- 
ovate, membranous margin, glabrous, or margin fimbrillate towards 
tip or with hairs on back. Calyx lobes about 3 mm long, broad, 
orbicular-ovate, obtuse, margin narrowly membranous. Corolla 
glabrous, with broadly ovate, short acuminate, about 1.5 mm long 
lobes. Capsule, ovoid-ellipsoidal, about 4 mm long, 4-locular. Seeds 
flattened, oblong-elliptic, dark colored, 2-3 mm long. June to August. 

On saline meadows and solonchaks.— European USSR: Upper 
Volga (rarely), Volga-Kama, Middle Dnieper, Volga-Don, Trans- Volga, 
Bessarabia, Black Sea Region, Crimea (?), Lower Don, Lower Volga. 
Caucasus: Ciscaucasia (rarely). Western Siberia: Ob’ Region (only in 
the south), Upper Tobol, Irtysh. Eastern Siberia: Lena-Kolyma (rarely), 
Angara-Sayan, Dauria. Soviet Central Asia: Aralo-Caspian, Balkhash 
Region. General distribution: Central Europe (the south), Mediterranean 
(western), Balkans-Asia Minor (Balkans), Dzhungaria-Kashgaria (?), 
Mongolia. Described from Montpellier. Type in London. 


Section 4. HoLopsytiium Pilger in Pflanzenr. H. 102, 101.— 
Capsule without transverse annular suture, indehiscent. Seeds large, 


143 


137 


with deeply grooved one side. Bracts and calyx lobes membranous, 
with extremely narrow midrib (keel). 


5. P. macrocarpa Cham. and Schlechtd. in Linnaea, I (1826) 166; 
Ldb. Fl. Ross. III, 483; Kom. xl. Kamch. III (1930) 95; Hulten, Fl. 
Kamtch. IV, 130; Pilger in Pflanzenr. H. 102, 101.—Ic.: Pilger l.c. f. 
14. 

Perennial. Plant 15—40 cm tall, with well-developed tap root and 
with 2-5 scapes, equaling or more or less exceeding leaves. Leaves 
thin, lanceolate or linear-lanceolate, acute, gradually narrowed at base 
into long and broad petiole, entire, glabrous. Scape glabrous in lower 
part, densely lanate upward, often subglabrous above in fruit. Spikes 
1.5-7 cm long, compact in flower (only lower flowers sometimes 
scattered) often lax in fruit. Bracts broad, ovate or orbicular, 3-4 mm 
long, membranous, equaling and enclosing calyx. Calyx lobes elliptical 
or broadly ovate, obtuse. Corolla lobes ovate or deltoid-ovate, 2—2.5 
mm long. Capsule elliptic, 5-7 mm long, without transverse annular 
suture along which upper part of capsules separates out, hence 
indehiscent, falling off as a whole, bilocular. Seeds 4-5 mm long, 
oblong-elliptic almost black. June to July. 

In marshy places and in wet meadows.—Soviet Far East: 
Kamchatka (Koragin Island, Commander Islands). General distribution: 
North America (Aleutian Islands, Alaska, Vancouver, Washington 
state). Described from Aleutian Islands (Unalaska). Cotype in 
Leningrad. 


Section 5. Coronopus DC. in Lam. and DC. FI. Frang. ed. 3, III 
(1805) 417, emend. Decne. in DC. Prodr. XIII, 1 (1852) 729.—Corolla 
tube pubescent. Ovules 3-5. Corolla persistent in fruit. Leaves mostly 
narrow, often pinnatidentate or pinnatisect. Annual or perennial 
herbaceous plant. 


6. P. coronopus L. Sp. pl. (1753) 115; Ldb. Fl. Ross. III, 480; O. 
and B. Fedch. Perech. rast. Turkest. V, 194; Grossh. Fl. Kavk. IV, 17; 
Pilger in Pflanzenr. H. 102, 126; Nikitin in Fl. Turkm. VI, 312.—P. 
filiformis C. Koch in Linnaea, XXI (1848) 709.—Ic.: Hegi. Ill. Fl. VI, 
1, f. 106; Pilger, l.c. f. 18-20; Nikitin, l.c. plate XL. 

Annual (rarely biennial ?). Plant (2)4—30(40) cm tall, with few or 
many, erect or ascending scapes, rarely with single scape. Leaves 
narrow, linear or lanceolate-linear, usually with few distant, narrow 
lateral lobes or teeth, rarely entire or subentire, somewhat densely 
hispid, rarely glabrous. Inflorescence (1)2—10(15) cm long, narrow, 
cylindrical, dense, flowers aggregated toward top. Bracts ovate, broadly 


144 


138 


membranous along margin, often fimbriate, short or fairly long 
acuminate, shorter than or almost equaling calyx. Calyx lobes 2-3 
mm long, glabrous or slightly pubescent, toward tip usually fimbrillate 
along margin; anterior calyx lobes elliptic, membranous along margin; 
posterior calyx lobes elliptic ovate, rarely keeled, somewhat winged 
along keel, wing fimbrillate along margin. Corolla lobes ovate or 
oblong-ovate, about 1 mm long. Capsule not exceeding calyx, 3-5 
seeded. Seed dark brown elliptic, lenticular, about 1 mm long. April 
to May. 

On sands, solonetzes, dry slopes, etc.— European USSR: Crimea. 
Caucasus: Dagestan, eastern Transcaucasia. Soviet Central Asia: Kara 
Kum (southern part), mountainous Turkmenia, Amu Darya, Pamiro- 
Alai (southern part). General distribution: Scandinavia (in the south), 
Central and Atlantic Europe, western and eastern Mediterranean, 
Balkans-Asia Minor, Armenia-Kurdistan, Iran. Described from western 
Europe. Type in London. 


7. P. maritima L. Sp. pl. (1753) 114; Ldb. Fl. Ross. III, 485; 
Schmalh. FI. II, 352; O. and B. Fedch. Perech. rast. Turkest. V, 197; 
Grossh. Fl. Kavk. IV, 17; Pilger in Pflanzenr. H. 102, 169.—P. salsa 
Pall. Reise, I, Anh. (1773) 486; M.B. fl. taur.-cauc. I, 110; Karyl. Fl. 
Zap. Sib. X, 2567.—P. schrenkii C. Koch in Linnaea, XXI (1848) 
710.—P. subpolaris Andrejew in Zhurn. Russk. bot. Obshch. XV, 4 
(1930) 298.—Ic.: Hegi, Ill. Fl. VI, 1, f. 107; Fi. Yugo-Vost. VI (1936) 
fig. 647 (sub P. salsa Pall.)—Exs.: GRF, No. 432; H.F.A.M. No. 174. 

Perennial. Plant (3)6—40(50) cm tall, with well-developed tap root, 
usually multiheaded, with few or many erect or ascending scapes. 
Leaves fleshy, linear or lanceolate-linear, usually canaliculate, subentire 
or remotely dentate, glabrous, rarely sparsely hairy. Scape densely 
upward appressed hairy, rarely subglabrous. Spikes (0.5)1—15 cm long, 
dense, narrowly cylindrical or capitate. Bracts lanceolate, ovate or 
broadly ovate, acute or obtuse, membranous along margin, sometimes 
with pointed keel, shorter than, equaling or longer (sometimes 
considerably longer) than calyx. Calyx lobes elliptic or ovate-elliptic, 
2—2.5(3) mm long, with well-developed straight keel, broadly 
membranous margin, glabrous or pubescent on back, fimbrillate along" 
margin. Corolla lobes ovate, 1—-1.5(2) mm long, glabrous or ciliolate 
along margin. Capsule (2)2.5—4 mm long, ovate or oblong-ovate, with 
1-3 seeds. Seeds oblong or ovate-oblong plano-convex, (1.5)2—2.5 
mm long. June to August (September). 

On solonchaks and saline meadows, occasionally on chalky and 
clayey outcrops in steppe and arid zones as well as along saline sea 
coast.—Arctic Region: Arctic Europe. European USSR: Karelia- 


145 


139 


Lapland, Dvina-Pechora, Baltic Region, Ladoga-IImen, Volga-Kama, 
Middle Dnieper, Volga-Don, Trans-Volga, Bessarabia, Black Sea 
Region, Crimea, Lower Don, Lower Volga. Caucasus: Ciscaucasia, 
Dagestan, southern Transcaucasia. Western Siberia: Upper Tobol, 
Irtysh. Eastern Siberia: Angara-Sayan, Dauriya. Soviet Central Asia: 
Aralo-Caspian, Balkhash Region, Tien Shan, rarely in mountainous 
Turkmenia. General distribution: Arctic Region, Scandinavia, Central 
and Atlantic Europe, western and eastern Mediterranean, Balkan-Asia 
Minor, Armenia-Kurdistan, Iran-Afghanistan (northern Iran), 
Dzhungaria-Kashgaria, Mongolia; moreover, in North and South 
America (Patagonia, Tierra del Fuego). Described from the coast of 
northern Europe. Type in London. 

Note. It is well known that P. maritima L. which covers a huge 
area, consisting of three distinct regions: Eurasian, North American 
and South American, exhibits considerable polymorphism. Its individual 
forms were often described as distinct species. Thus, certain authors 
consider P. salsa Pall. a distinct species, a form widespread in the 
Southern European part of the USSR, in southern Siberia, Kazakhstan 
and Mongolia. The main distinctive feature of this form is short-ciliate 
corolla lobes. It is necessary to observe that within the limits of the. 
indicated range of this form, one often comes across specimens of P. 
maritima with corolla lobes completely lacking cilia. Moreover, certain 
specimens, seen by the present authors in Kazakhstan and, especially, 
in Mongolia are so similar to specimens from the Baltic Sea Coast, 
that their placement in the same species. and even in the same form 
could not be questioned. Thus, segregation of P. salsa Pall. as a distinct 
species is without basis. It makes more sense to segregate the dwarf 
form of P. maritima, which has short, few-flowered, capitate 
inflorescences (P. maritima L. var. glauca Hornem. = P. schrenkii C. 
Koch), distributed mainly along the coast of the Arctic Ocean. However, 
this form is connected with the ‘typical’ P. maritima L. from the 
banks of the Baltic and White seas by so many transitional forms that 
it is impossible to consider it as a distinct species. Plants transitional 
between typical P. maritima L. and L. schrenkii C. Koch, were 
described by V.N. Andreev as a distinct species.—P. subpolaris 
Andrejew. The author made a number of mistakes describing it. For 
example, the original diagnosis indicates: “Petala 3-4 mm lg.”, when, 
in fact, the corolla lobes of an authentic specimen never exceed 2 mm. 


Section 6. MESEMBRYNIA Decne. in DC. Prodr. XIII, 1(1852) 705— 
Ovules 5, 1 locule of ovary with 2 ovules, the remaining with 3. 
Anterior calyx lobes different from posterior. Corolla lobes often 
reflexed. Annual or biennial herbaceous plant. 


146 


140 


8. P. schwarzenbergiana Schur. in Verh. Sieb. Ver. Nat. VI (1855) 
3, X (1859) 79; Pilger in Pflanzenr. H. 102, 271; Bilyk in Bot. zhurn. 
Akad. nauk URSR, VIII, 1, 95. 

Annual or perennial. Plant 12-25 cm tall with one or often several 
(5-8), ascending, rarely erect scapes, usually much longer than leaves. 
Leaves thick, oblong-lanceolate or ovate-lanceolate, gradually narrowed 
at base into a long petiole, acute or subobtuse, margin subentire or 
dentate, usually distinctly 3-veined (rarely having 2 more less distinct 
lateral veins). Scapes glabrous or sparsely appressed hairy. Spikes 3-— 
9 cm long, slender, dense, sometimes slightly interrupted only at base. 
Bracts about 2 mm long, elliptic, obtuse, membranous along margin, 
equaling calyx lobes or somewhat shorter. Calyx lobes, 2—-2.5 mm 
long, elliptic, membranous, 2 anterior ones with distinct herbaceous 
keel, posterior ones with barely noticeable keel. Corolla with ovate or 
oblong-ovate lobes, about 1.5 mm long. Capsule cuneate-ovate, much 
longer than calyx, 4-5 seeded. Seeds about 1 mm long, irregularly 
elliptic. June. 

On saline meadows.— European USSR: Black Sea Region (reported 
from Bashtan District of Nikolaev Region). General distribution: 
Central Europe (Hungary, Romania). Described from Hungary. Type 
probably in Paris. 


9. P. depressa Willd. Enum. pl. Hort. Bot. Berol. Suppl. (1813) 
8; Ganeschin in Tr. Bot. myz. AN, XIII, 193; Pilger in Pflanzenr. H. 
102, 272; Kryl. Fl. Zap. Sib. X, 2559.—P. sibirica Poir. Encycl. meth. 
Suppl. IV (1816) 433; Ldb. Fl. Ross. III, 478.—P. leptostachys Ldb. 
FI. Ross. III (1847-1849) 479.—P. paludosa Turcz. ex Decne. in DC. 
Prodr. XIII, 1 (1852) 695.—Ie.: Pilger, l.c. f. 29. 

Annual or biennial. Plant 5—40(65) cm tall, with 1 or often several 
(2-15), erect, ascending scapes. Leaves oblong-elliptic, elliptic or 
ovate-lanceolate, narrowed at base into a petiole, acute or obtuse, 
entire or remotely dentate, glabrous or slightly appressed hairy, 
distinctly 5—7-veined, petiole shorter than, rarely equaling or longer 
than lamina. Scapes sulcate, sparsely appressed hairy. Inflorescence 
3-15(20) cm long, slender and dense, usually interrupted in lower 
part. Bracts ovate, acute or subacute, membranous along margin,. 
equaling calyx lobes or somewhat shorter. Calyx lobes broadly obovate 
or elliptic, about 2 mm long, membranous, but all 5 calyx lobes with 
distinct herbaceous keel. Corolla lobes ovate or oblong-ovate, about 1 
mm long. Capsule ovate or ovate-cuneate, about 3 mm long, with (4)5 
seeds. Seeds elliptic-oblong, about 1.5 mm long, almost black. June to 
August. 


147 


141 


In meadows, coastal sand and slopes, and also fields, roadsides, 
pastures, etc.— Western Siberia: Ob’ Region (in the south), Upper 
Tobol (only in north), Irtysh, Altai. Eastern Siberia: Yenisei, Lena- 
Kolyma, Angara-Sayan, Dauria. Soviet Far East: Zeya-Bureya, Uda 
River area, Ussuri, Sakhalin. Soviet Central Asia: Dzhungaria- 
Tarbagatai, Tien Shan. General distribution: India-Himalayas, 
Dzhungaria-Kashgaria, Mongolia, Japan, China, Tibet. Described from 
cultivated specimens, grown in Berlin Botanical Garden. Type in Berlin. 


10. P. camtschatica Link, Enum. pl. Hort. Bot. Berol. Abt. I 
(1821) 120; Cham. and Schlechtd. in Linnaea, I, 165; Pilger in 
Pflanzenr. H. 102, 274.—P. kamtschatica Ldb. FI. Ross. III, I (1847- 
1849) 478; Kom. Fl. Kamch. III, 94,—P. arctica Decne. in DC. Prodr. 
XIII, I (1852) 700. 

Perennial. Plant 5—35 cm tall, with several (2-12) usually ascending 
scapes, exceeding leaves. Leaves thicker, oblong-elliptic, elliptic or 
oblong-ovate, narrowed at base into petiole, obtuse or subacute, entire 
or somewhat dentate along margin, usually 5-veined, young leaves 
densely mature leaves more sparsely white hairy, petiole usually shorter 
than lamina, rarely as long or longer. Scapes sulcate, densely hairy, 
young occasionally white due to dense pubescence. Spike 2-12 cm 
long, dense or with several scattered lower flowers. Bracts ovate or 
ovate-lanceolate, membranous along margin, keeled, equaling calyx 
lobes or shorter. Calyx lobes elliptic or obovate, 2.5-3 mm long, 
margin broadly membranous, but with well-developed herbaceous keel. 
Corolla lobes fairly broad, ovate, 1-1.5 mm long. Capsule ovate or 
ovate-cuneate, 3-4 mm long, 4—5 seeded. Seeds elliptic or oblong- 
elliptic, 1.5—2 mm long, black. May to August (September). 

On sands, argillaceous soils and rocks (sometimes in weedy places) 
mainly along seacoasts.— Soviet Far East: Kamchatka, Uda River area, 
Ussuri, Sakhalin. General distribution: Japan, China. Described from 
cultivated specimens, grown in Berlin Botanical Garden. Type in Berlin. 


11. P. arachnoidea Schrenk in Fisch. and Mey. Enum. pl. nov. 
(1841) 16; Ldb. Fl. Ross. III, 484; O. and B. Fedch. Perech. rast. 
Turkest. V, 195; Pilger in Pflanzenr. H. 102, 276. 

Perennial. Plant 3-15(25) cm tall, with thick, vertical, often 
multiheaded root and with 3-10(20) ascending or erect scapes much 
longer than leaves, rarely shorter. Leaves narrowly lanceolate, 
lanceolate or elliptic, acute, gradually narrowed at base into petiole, 
usually short and broad, entire or dentate; young leaves (as well as 
scape) densely arachnoid-lanate, mature more rarely so. Spikes 0.5— 
4(5) cm long, cylindrical, dense, sometimes interrupted in lower part, 


148 


142 


with more or less scattered flowers. Bracts ovate or broadly ovate, 
margin broadly membranous, obtuse, equaling calyx lobes or somewhat 
shorter. Calyx lobes 2-3 mm long, elliptic or ovate-elliptic, margin 
broad, membranous, all lobes with distinct herbaceous keel, sometimes 
lanate along margin. Corolla lobes broad, ovate or orbicular-ovate, 1— 
1.5 mm long. Capsule ovoid or oblong-ovoid, 3—4 mm long, with 2- 
3(4) seeds. Seeds relatively larger, 2-3 mm long, elliptic-oblong, or 
ovate-oblong, black. June to August (September) (Plate VII, fig. 3). 

In clayey soils, rubbly and stony slopes in high-mountain regions.— 
Soviet Central Asia: Dzhungaria-Tarbagatai, Tien Shan, Pamiro-Alai 
(Eastern Pamir, Alai and Turkestan ranges). General distribution: 
Dzhungaria-Kashgaria. Described from Tarbagatai. Type in Leningrad. 


12. P. komarovii Pavl. in Byull. Mosk. Obshch. ispyt. prir. 
XXXVIII, issue 1-2 (1929) 130.—Ic.: Pavlov. l.c. fig. 8. 

Perennial. Plant 3-8 cm tall, with well-developed tap root and 
with several procumbent, ascending or almost erect scapes. Leaves 
thicker, linear or linear-lanceolate, (2)3—6 mm broad, entire, glabrous 
or sparsely hairy above. Scapes sparsely appressed long hairy. Spikes 
fairly dense, capitate or short cylindrical, 4-10(12) mm long, few- 
flowered. Bracts ovate or deltoid-ovate, keeled, herbaceous, 
membranous only along margin, equaling calyx or slightly longer. 
Calyx lobes elliptic or ovate-elliptic, 1.5-2 mm long, membranous, 
but with well-developed herbaceous keel. Corolla lobes about 1 mm 
long. Capsule ovate, 2-3 mm long, 4-seeded. Seed ovoid-oblong, 0.5— 
2 mm long, nearly black. May to June. 

In high mountain meadows and on damp slopes.—Soviet Central 
Asia: Dzhungaria-Tarbagatai (Saur range). General distribution: 
Mongolia. Described from Mongolia (Khanghai). Type in Leningrad. 


Section 7. LAMPROSANTHA Decne. In DC. Prodr. XIII, 1 (1852) 
697.—Ovules 4(6). Corolla lobes whitish, lustrous. Rather large 
perennial plants, usually with well-developed tap root. Spike more or 
less dense, many-flowered. 


13. P. media L. Sp. pl. (1753) 113; M.B. Fl. taur.-cauc. I, 109; 
Ldb. Fl. Ross. III, 480; Schmalh. Fl. II, 351; O. and B. Fedch. rast. 
Turkest. V, 194; Grossh. Fl. Kavk. IV, 17; Pilger in Pflanzenr. H. 
102, 277; Kryl. Fl. Zap. Sib. X, 2563.—Ic.: Hegi, Ill. Fl. VI, I, tab. 
246, f. 4.—Exs.: Pl. Finl. exs. No. 247. 

Perennial. Plant 15-70 cm tall, with several scapes, usually 
ascending at base, erect above. Leaves elliptic or ovate, rarely ovate- 
lanceolate, usually no more than 2-2 times as long as broad, generally 


151 


143 


pubescent (especially beneath). Subentire or with few remote teeth 


along margin, usually acute, narrowed at base into short (3-10 times 


as short as lamina) and broad petiole, or subsessile; however, in shaded 
forms, petiole longer, sometimes only slightly shorter than or even 
equaling lamina. Scapes pubescent, upward appressed hairy. 
Inflorescence dense, cylindrical, 2—6(8) cm long. Bracts ovate or 
oblong-ovate, subobtuse, membranous along margin, equal to or almost 
equaling calyx. Calyx lobes 1.5—2 mm long, elliptical or ovate-elliptical, 
membranous, but all with distinct herbaceous keel. Corolla lobes ovate 
or oblong-ovate, 1.5—2 mm long. Capsule ovoid, 2—2.5(3) mm long, 
with 2—5(6) seeds. (May) June to August (Plate VII, fig. 1). 

In floodplain, dry-valley and high-mountain meadows and in open 
forests as well as fallow land, roadsides, etc., mainly in forest plain 
zone.— European USSR: All regions. Western Siberia: All regions. 
Eastern Siberia: All regions. Soviet Far East: Kamchatka (rarely as 
introduced). Soviet Central Asia: Dzhungaria-Tarbagatai, as far as Tien 
Shan. General distribution: Scandinavia, Central and Atlantic Europe, 
western Mediterranean, Balkans-Asia Minor, extending up to northern 
Iran. Described from Western Europe. Type in London. 


14. P. stepposa Kupr. in Tr. Bot. inst. AN SSSR, ser. 1; vyp. 
3(1936) 280; Kryl. Fl. Zap. Sib. X, 2564.—P. media var. urvilleana 
Rapin in Mém. Soc. Linn. Paris, VI (1827) 453; O. and B. Fedch. 
Perech. rast. Turkest. V, 194; Grossh. Fl. Kavk. IV, 17; Pilger in 
Pflanzenr. H. 102, 279.—Ic.: Kupriyanova, l|.c. fig. 1.—Exs.: GRF, 
No. 386 (sub P. media L.). 

Perennial. Plant 20—70 cm tall, with distinct tap root and several 
scapes, erect or slightly ascending at base. Leaves elliptic (rarely 
elliptic-ovate), lanceolate or narrowly lanceolate, 2¥2—5 times as long 
as broad, gradually narrowed toward base and apex, more or less 
hairy (especially beneath), with petioles almost equaling lamina or 
shorter, but not more than 2-3 times as long, margin entire or subentire. 
Scapes somewhat densely appressed hairy upward. Inflorescence dense, 
cylindrical, (4)6-12(20) cm long. Bracts ovate or oblong-deltoid, 
subacute, keeled, membranous along margin, nearly equaling calyx. 
Calyx lobes elliptic or ovate-elliptic, 2-2.5 mm long, membranous, 
but with distinct herbaceous keel. Corolla lobes oblong-lanceolate or 
oblong-ovate, 2-2.5 mm long. Capsule ovate, 2.5—-3 mm long, with 2— 
5(6) seeds. May to July (August) (Plate VII). 

Steppes, saline meadows and fields, occasionally in forest edges.— 
European USSR: Volga-Kama, Middle Dnieper, Volga-Don, Trans- 
Volga Region, Bessarabia, Black Sea Region, Crimea, Lower Don, 
Lower Volga. Caucasus: Ciscaucasia, Dagestan, eastern and western 


144 


Plate VII. 
1. Plantago media L.—2. P. lagocephala Bge.—3. P. arachnoidea Schrenk. 


149 


152 


145 


Transcaucasia. Western Siberia: Upper Tobol, Irtysh, Altai. Eastern 
Siberia: Angara-Sayan. Soviet Central Asia: Dzhungaria-Tarbagatai, 
Tien Shan. General distribution: Central Europe (Hungary). Described 
(under the name P. media var. urvilleana Rapin.) from Crimea (Kerch 
area). Type in Paris. 


15: P. maxima Juss. ex Jacq. Collect. ad Bot. I (1786) 82; An. 
Hort. Kew. I, 151; Ldb. Fl. Ross. II, 480; Schmalh. FI. II, 350; O. and 
B. Fedch. Perech. rast. Turkest. V, 194, Grossh. Fl. Kavk. IV, 17; 
Pilger in Pflanzenr. H. 102, 281; Kryl. Fl. Zap. Sib. X, 2561.—Ic.: 
Jacq. Ic. pl. rar. I, tab. 26. 

Perennial. Plant 20—70(90) cm tall, with thick tap root and 1 or 
several (2-9), usually erect shapes. Leaves thicker, short, ovate or 
orbicular-ovate, rarely elliptic, short acuminate or obtuse, at base short 
and broadly cuneate, entire or denticulate, 7—11-veined, long petiolate, 
pubescent on both surfaces or only beneath, along with petioles, usually 
blackened when dry. Petioles semicircular in cross section, sulcate at 
top, longer than lamina, rarely as long or shorter. Scape densely 
appressed hairy above. Spikes, compact, or fairly thick, 5—20 cm long, 
with closely-packed flowers. Bracts linear-oblong or oblong-ovate, 
margin membranous, somewhat shorter than calyx lobes. Calyx lobes 
2.5—3 mm long, elliptic or ovate-elliptic, membranous but with distinct 
keel. Corolla lobes oblong-ovate, acuminate, about 2 mm long, silver 
white. Capsule oblong-elliptic, about 3 mm long, 4-seeded. Seeds 
flattened, oblong-elliptic, about 2 mm long. June to August. 

In meadows, usually saline ones and also mixed-grass steppes.— 
European USSR: Volga-Kama, Upper Dnieper, Middle Dnieper, Volga- 
Don, Trans-Volga, Black Sea Region, Lower Don. Caucasus: 
Ciscaucasia. Western Siberia: Ob’ Region (in south), Upper Tobol, 
Irtysh. Eastern Siberia: Angara-Sayan. Soviet Central Asia: Aralo- 
Caspian, as far as Balkhash Region. General distribution: Central 
Europe (in south), extending as far as Dzhungaria-Kashgaria 
(Dzhungaria, Chernyi Irtysh valley). Described from Siberia. Type in 
Paris. 


16. P. canescens Adams in Mém. Soc. Nat. Mosc. IX (1834) 233; 
Ldb. FI. Ross. III, 483; Pilger in Pflanzenr. H. 102, 282.—Ic.: Adams, 
l.c. tab. XIII, f.1. 

Perennial. Plant (8)10—30 cm tall, with distinct tap root, sometimes 
multiheaded, and with 1 or several (2—8) scapes, usually much longer 
than leaves. Leaves lanceolate or linear-lanceolate, 0.5—1.5(2) cm broad, 
(3)5-veined, gradually narrowed toward tip, subacute, narrowed at base 
into short or long petiole, more or less densely or sparsely ianate, 
rarely subglabrous. Scapes, especially young ones, more or less, hairy 


155 


146 


(upward appressed hairy), rarely subglabrous. Spikes 2—5(6) cm long, 
fairly dense, occasionally somewhat interrupted at base, with scattered 
flowers. Bracts ovate or broadly ovate, margin membranous, equaling 
calyx lobes or slightly shorter. Calyx lobes elliptic or broadly ovate, 
2—2.5 mm long, membranous, but all with distinct herbaceous keel. 
Corolla lobes ovate or oblong-ovate, 1.5-2 mm long, acute. Capsule 
ovoid, about 3 mm long, with (2)4—5 seeds. Seed flattened, oblong- 
elliptic, about 2 mm long, almost black. May to July. 

On dry meadows and slopes in scrub.—Eastern Siberia: Lena- 
Kolyma. General distribution: North America. Described from Yakutsk 
area. Type unknown. 


Section 8. OREADES Decne. in DC. Prodr. XIII, 1 (1852) 717, emend. 
Pilger in Fedde, Repert. sp. n. XXIII (1926) 241.—Ovules 2. Bracts 
broad, enclosing calyx. Calyx lobes membranous, with weakly 
developed midrib (keel). Perennial, high-altitude herbs. 


17. P. atrata Hoppe, Bot. Taschenb. (1799) 85; Pilger in Pflanzenr. 
H. 102, 281.—P. montana auct. non Huds. (1762); Lam. Ill. gen. 
(1791) 341; Popov, Ocherk rastit. i flory Karpat (1949) 235.—P. 
saxatilis M.B. Fl. taur.-cauc. 1 (1808) 109; Ldb. Fl. Ross. III, 483; 
Grossh. Fl. Kavk. IV, 18.—P. montana var. saxatilis Schmalh. FI. I 
(1897) 351.—P. caucasica Popova in Soodshch. Akad. nauk. Gruz. 
SSR, IX, Nos. 8-9 (1948) 591.—Ic.: Pilger, l.c. f. 30; Popova, l.c. 
593: 

Perennial. Plant 5—30 cm tall, with pronounced tap root, sometimes 
multiheaded, and usually with several (2-10) erect or ascending scapes. 
Leaves linear or lanceolate, 3-15 mm broad, gradually narrowed toward 
base and apex, acute, entire or dentate, as in scape densely appressed 
hairy or glabrous. Inflorescence dense, capitate or short cylindrical, 
0.8-3(4) cm long. Bracts enclosing calyx, broad (breadth seldom 
exceeds length), variable, irregularly shaped, ovate, broadly elliptic or 
suborbicular, (3.5)4-6(7) mm long, membranous, but with distinct 
herbaceous keel, brown or whitish, glabrous or, more or less hairy 
along keel and margin (or only along margin). Calyx lobes shorter 
than bracts, elliptic or orbicular-elliptic, 2.5—4 mm long, almost wholly 
membranous, sometimes with long hairs along margin. Corolla lobes 
ovate or oblong-ovate, 2-3 mm long. Capsule oblong-ovate, 4—6(7) 
mm long, 2-seeded. Seeds ovoid or oblong-ovoid, about 4 mm long, 
sulcate. May to July. 

In subalpine and alpine zones in the upper part of mid-altitudinal 
mountain zone in meadows and on rubbly slopes.— European USSR: 


156 


147 


Upper Dniester, Crimea (?). Caucasus: All regions. General 
distribution: Central Europe (southern part), western Mediterranean, 
Balkans-Asia Minor, Armenia-Kurdistan, Iran. Described from Salzburg 
area. Cotype in Leningrad. 


Section 9. GENTIANOIWES Pilger in Pflanzenr. H. 102 (1937) 306.— 
Ovules 3-7. Bracts and calyx lobes broad, broadly membranous along 
margin, with distinct herbaceous keel. Perennial, herbaceous plants 
found at high altitudes. 


18. P. gentianoides Sibth. et Smith, Fl. graec. Prodr. I (1806) 
101; Boiss. Fl. or. IV, 879; Pilger in Pflanzenr. H. 102, 306.—P. 
tatarica Decne. in DC. Prodr. XIII, 1 (1852) 696.—P. griffithii Decne. 
l.c. 700; Boiss. Fl. or. IV, 880; O. and B. Fedch. Perech. rast. Turkest. 
V, 195.—P. sogdiana M. Pop. in Rast. zapov. Guralash i Zaam. lesnoi 
dachi (1937) 35. 

Perennial. Plant 5—20 cm tall, with several or many (up to 15-20), 
erect or ascending, scapes. Leaves ovate-lanceolate, ovate or elliptic, 
acute or subobtuse, gradually narrowed from broad base into short 
and broad or fairly long petioles, entire or sparsely dentate, glabrous 
like scapes, rarely somewhat densely pubescent. Spikes subcapitate or 
cylindrical, 0.7—-3(4) cm long, thick and compact or lax. Bracts ovate 
or ovate-deltoid, broadly membranous along margin, equaling calyx 
lobes or somewhat shorter. Calyx lobes about 2 mm long, elliptic or 
ovate-elliptic, membranous, all with distinct herbaceous keel. Corolla 
lobes ovate or ovate-elliptic, about 1.5 mm long. Capsule ovoid, 
ellipsoid or subglobse, about 3 mm long, with (3)4—7 seeds. Seeds 
flattened, outline unevenly ovate or elliptic, 1.5-2.5 mm long. June to 
August. 

In wet marshy meadows, primarily in high-mountain region.— 
Soviet Central Asia: Pamiro-Alai (Pamir, Zeravshan Basin, Turkestan 
Range, rarely on southern slopes of Hissar Range). General distribution: 
Balkans-Asia Minor, Iran, India-Himalayas. Described from Greece. 
Type in Oxford. 


Section 10. ARNoGLossuM Decne. in DC. Prodr. XIII, 1 (1852) 
714.—Capsule 2-seeded. Seed with 1 side convex and the other sulcate. 
2 anterior calyx lobes connate, posterior calyx lobes free. Spike dense, 
with closely-packed flowers. Scape more or less distinctly 
longitudinally sulcate. Annual or perennial, herbaceous plants with 
lanceolate leaves. 


148 


Plate VIII. 
Plantago stepposa Kuprt. 


153 


157 


149 


19. P. lanceolata L. Sp. pl. (1753) 113; M.B. fl. taur.-cauc. I, 
109; Ldb. Fl. Ross. III, 481; Schmalh. Fl. II, 351; O. and B. Fedch. 
Perech. rast. Turkest. V, 195; Grossh. Fl. Kavk. IV, 19; Pilger in 
Pflanzenr. H. 102, 313; Kryl. Fl. Zap. Sib. X, 2565.—P. lanuginosa 
Bastard, Essai Fl. Dep. Maine and Loire (1809) 160; Grossh. I.c. 18; 
Heideman. Opred rast. Moldavsk SSR, 225.—P. glauca C.A.M. 
Verzeichn. Pfl. Cauc. (1831) 115; Ldb. l.c. 483.—P. fontiscurvae Kom. 
in Kom and Alis. Opred. rast. Dalnevost. kr. II (1932) 942;—P. 
altissima auct. non L.; Grossh. Fl. Kavk. IV (1934) 18.—P. glabriflora 
Sakalo in Bot. zhurn. Akad. nauk. URSR, IV, No. 3-4 (1948) 84.— 
Ic.: Hegi, Ill. Fl. VI, tab. 246; Kom. and Allis. l.c. tabl. 281; Pilger, 
l.c. f. 32; Nikitin, in Fl. Turkm. VI, plate XLI. 

Perennial. Plant 8-70 cm tall, with well-developed tap root and 
with several or numerous (25-30) erect or ascending scapes. Leaves 
elliptic-lanceolate, lanceolate or linear-lanceolate, 3—5(7)-veined, 
gradually narrowed toward base and apex, acute, with short or fairly 
long petioles; entire, rarely sparsely dentate, glabrous or densely hairy 
or lanate. Scapes distinctly longitudinally sulcate, usually pubescent, 
upward appressed hairy. Spikes dense, compact, capitate or cylindrical, 
0.5—7 cm long. Bracts ovate or elliptic, membranous, with distinct 
herbaceous keel not reaching the tip of bracts, short or long acuminate, 
equaling or exceeding calyx, glabrous or sparsely hairy. Calyx lobes 
2.5-3.5 mm long, membranous, all with distinct herbaceous keel, 
glabrous, rarely more or less hairy along back, 2 anterior calyx lobes 
connate, forming a broad, shortly bilobed scale with 2 keels. Corolla 
lobes ovate or oblong-ovate, acuminate, about 1.2 mm long. Capsule 
elliptic, about 3 mm long, 2-seeded. Seeds oblong or oblong-elliptic, 
with one side convex, the other sulcate. April to September (October). 

Dry meadows, wastelands, roadsides, banks of river and irrigation 
ditches, fallow land, open dry slopes, etc., usually on plains as well 
as in mountains, reaching to high mountain zones.— European USSR: 
All regions. Caucasus: All regions. Western Siberia: Upper Tobol. 
Soviet Central Asia: All regions. Occasionally in Eastern Siberia and 
in Far Eastern regions as introduced plant. General distribution: 
Scandinavia and Atlantic Europe, western and eastern Mediterranean, 
Balkan States-Asia Minor, Armenia-Kurdistan, Iran, India-Himalayas, 
Dzhungaria-Kashgaria, moreover, as an adventive plant in many other 
countries (Abyssinia, North and South America, etc.). Described from 
Western Europe. Type in London. 

Note 1. Under drought conditions, the same plant of P. lanceolata 
may have two types of leaves: glabrous and broad, developing in 
spring, and more or less hairy and narrower, developing in the 
beginning of summer. 


158 


150 


Note 2. Certain authors report from the territory of the USSR, in 
particular, the Caucasus, two other species of section Arnoglossum, 
namely P. altissima L. and P. lagopus L. In fact, so far these species 
have not been found in the USSR. 


Section II]. LEucopsyLLiuM Decne. in DC. Prodr. XIII, 1 (1852) 
704.—Ovules 2. Capsule usually 2-seeded. Seeds with one side deeply 
concave or sulcate. Spike mostly short and dense. Small annual or 
perennial plants rarely semishrubs. Leaves narrow, often linear or 
linear-lanceolate. 


20. P. ovata Forsk. Fl. Aegipt. Arab. (1775) 31; Grossh. Fl. Kavk. 


IV, 19; Pilger and Pflanzenr. H. 102, 347; Nikitin in fl. Turkm. VI, 


316.—P. villosa Moench. Meth. Pl. (1794) 459; Ldb. Fl. Ross. III, 
484; O. and B. Fedch. Perech. rast. Turkest. V, 196.—Ic.: Pilger, l.c. 
f. 35; Nikitin, l.c. plate XLI, fig. 2.—Exs.: Sintenis. It. transc.-pers. 
No. 1867. 

Annual (sometimes biennial?). Plant 3—20 cm tall, with several or 
many, ascending, rarely erect scapes. Sometimes several leaves in 
rosettes, lateral rosettes on shortened (all clearly noticeable) lateral 
shoots. Leaves linear, (1)2-6(9) mm broad, entire or remotely 
denticulate, more or less densely, rarely sparsely, lanate. Scapes shorter 
than leaves or somewhat longer, more or less lanate. Inflorescence 
dense, elliptic or short cylindrical, 1-3 cm long. Bracts ovate-orbicular 
or orbicular, equaling calyx or slightly shorter, membranous, with 
distinct herbaceous keel, glabrous, sometimes fimbrillate along margin. 
Calyx lobes obovate or elliptic, about 3 mm long, membranous, all 
with distinct herbaceous keel, glabrous. Corolla lobes orbicular-ovate 
or orbicular, about 2.5 mm long, short acuminate or subacute. Capsule 
ovoid-ellipsoidal, about 3 mm long. Seeds oblong-ovate or oblong- 
elliptic, 2-2.5 mm long, with one side convex, the other deeply 
concave. April to May. 

In sands or takyrs [clay soils in deserts] and dry rubbly slopes.— 
Caucasus: Eastern Transcaucasia. Soviet Central Asia: mountainous 
Turkmenia and, partially, Kara Kum. General distribution: Western 
and eastern Mediterranean, Armenia-Kurdistan, Iran, India-Himalayas. 
Described from Egypt (Alexandria). Type, probably, in Kiel. 

Economic importance. The seeds of P. ovata, in the Indian 
pharmacopoeia, are used with great success for treating acute gastric 
disorders. In the USSR, some experiments in the culture of P. ovata 
have been done (Sovetskie subtropiki, No. 11 (74) 1940). 


159 


151 


21. P. minuta Pall. Reise, III (1776) Anhang 716; M.B. FI. taur.- 
cauc. I (1808) 109; Ldb. Fl. Ross. III, 484; Schmalh. Fl. II, 352; O. 
and B. Fedch. Perech. rast. Turkest. V, 196; Grossh. Fl. Kavk. IV, 19; 
Pilger in Pflanzenr. H. 102, 351; Kryl. Fl. Zap. Sib. X, 2566.—P. 
lessingii Fisch. and Mey. in Ind. II Sem. Horti Petrop. (1835) 47; Ldb. 
l.c. 485.—P. latibracteata Trautv. Pl. Imag. and Descr. Fl. Russ. illustr. 
(1844) 20.—P. mongolica Decne. in DC. Prodr. XIII, 1 (1852) 707.— 
Ic.: Pall. l.c. tab. E, f. 1; Trautv. l.c. tab. 14. 

Annual. Plant (1.5)2—12 cm tall, or often with numerous spreading 
or ascending scapes. Leaves linear or linear-lanceolate, (1)2—-8 mm 
broad, densely or diffusely lanate, sometimes only beneath, rarely 
subglabrous. Scape more or less densely hairy. Spike capitate or 
cylindrical, rarely lax. Bracts broad, ovate or orbicular-ovate (their 
breadth rarely exceeding length), usually much shorter than calyx, 
rarely almost as long, membranous, but with distinct herbaceous keel, 
glabrous or more or less densely short- or long hairy. Calyx lobes 
elliptic-oblong, elliptic or broadly elliptic, about 3 mm long, 
membranous, with distinct keel, glabrous or more or less densely hairy. 
Corolla lobes oblong-ovate or oblong-elliptic, about 1.5 mm long. 
Capsule ovoid-cuneate, 3.5—5 mm long, much longer than calyx. Seeds 
oblong or oblong-elliptic, 3-4 mm long, with one side convex, the 
other deeply concave. May to June (August). 

In Salsola and Artemisia deserts, in clayey and rubbly soils, and 
also in takyrs [clay soils in deserts], shingles and chalk outcrops, 
etc.— European USSR: Lower Volga. Caucasus: Eastern and southern 
Transcaucasia. Soviet Central Asia: Aralo-Caspian, Balkhash Region, 
Tien Shan. General distribution: Dzhungaria-Kashgaria, Mongolia, 
extends to Tibet. Described from western Kazakhstan. Type in London. 

Note. The species is highly variable in the degree of the pubescence 
of the bracts and calyx segments, as well as the leaves. A form with 
more or less hairy bracts and calyx lobes is often regarded as a distinct 
species—P. lessingii Fisch. et Mey. (or P. mongolica Decne.). This 
species can hardly be distinguished from “typical” P. minuta Pall., 
since the glabrous (i.e., “typical’) and hairy forms are connected by 
intermediates. The geographical ranges of these forms nearly coincide. 
However, in Trans-Volga, the hairy form (with more or less hairy bracts 
and calyx lobes) evidently is not found, whereas in Soviet Central Asia 
the hairy form is predominant and widespread, and sometimes the 
pubescence is considerably more developed than in plants from the 
more western parts of the range of the species. The most densely hairy 
specimens from Soviet Central Asia were described under the name P. 
mongolica Decne. 


160 


152 


22. P. loeflingii L. Sp. pl. (1753) 115; Ldb. Fl. Ross. III, 485; 
Grossh. Fl. Kavk. IV, 19; Pilger in Pflanzenr. H. 102, 352.—Ic.: Pilger, 
l.c. f. 35; Karyagin, Fl. Apsherona, Plate 18. 

Annual. Plant 2-15 cm tall, with several or numerous, usually 
ascending, rarely erect scapes. Leaves narrowly linear or linear- 
lanceolate, (0.5)1.5—7 mm broad, entire or remotely denticulate, and 
like scape, more or less hairy. Spikes capitate or short cylindrical, 
dense, 0.5—2(2.5) cm long. Bracts broad, enclosing calyx, transversely 
elliptic (breadth exceeding length), membranous, but with thickened 
herbaceous keel continued into obtuse apex projecting above margin 
of scale, usually glabrous. Calyx lobes ovate-orbicular, about 2 mm 
long, almost whoHy membranous (only at base with distinct keel), 
glabrous. Corolla lobes ovate-lanceolate or lanceolate, acute only 1 
mm long. Capsule cuneate-ovate, about 3 mm long, often depressed at 
tip. Seeds elliptic-oblong, about 2 mm long, with one side convex, the 
other sulcate. April to May. 

On littoral sands, limestone, clayey slopes, etc.— Caucasus: Eastern 
and southern Transcaucasia (western Transcaucasia?). General 
distribution: Portugal, western and eastern Mediterranean, Armenia- 
Kurdistan, Iran. Described from Spain. Type in London: 

Economic importance. There is some information that this species 
is used in Iran as medicine. 


23. P. notata Lagasca, Gen. and sp. pl. (1816) 7; Grossh. FI. 
Kavk. IV, 19; Pilger in Pflanzenr. H. 102, 354.—P. syrtica Viviani. 
Fl. Lyb. sp. (1824) 7.—P. praecox C.A.M. Verzeichn. Pfl. Cauc. (1831) 
115; Ldb. Fl. Ross. III, 484; Karyagin, Fl. Apsherona, 349,—Ic.: 
Lagasca, l.c. tab III, f. 2; Pilger, l.c. f. 35; Karyagin, l.c. plate 18, 
fig. 6. 

Annual. Plant 3-10 cm tall, with several or numerous usually 
ascending, rarely erect scapes. Leaves linear-lanceolate or lanceolate, 
1-8 mm broad, gradually narrowed toward base and apex, with several 
fairly large teeth along margin, more or less lanate. Scapes more or 
less densely pubescent with reddish hairs. Spikes dense, ovoid or short 
cylindrical, 1-2.5(3) cm long, densely lanate (especially the young). 
Bracts transversely elliptical (breadth exceeding length), longer than 
calyx, membranous, with distinct herbaceous keel, lanate along keel. 
Calyx lobes orbicular-elliptic, about 2 mm long, wholly membranous, 
with a barely noticeable keel only at base, ciliate along margin. Corolla 
lobes orbicular-ovate, about 2 mm long, obtuse or subobtuse. Capsule 
ellipsoidal-orbicular, about 2.5 mm long. Seeds oblong-elliptic, about 
2 mm long, with one side convex, the other deeply grooved. April to 
May. 


161 


153 


On littoral sands, dry slopes and weedy places.— Caucasus: Eastern 
Transcaucasia (Apsheron). General distribution: Western and eastern 
Mediterranean, Armenia-Kurdistan, in parts of Iran. Described from 
Spain. Type in Madrid (?). 


24. P. lachnantha Bge. in Mém. sav. étrang. Acad. Sc. Pétersb. 
VII (1851) 446; O. and B. Fedch. Perech. rast. Turkest. II, 197; Pilger 
in Pflanzenr. H. 102, 355; Nikitin in Fl. Turkm. VI, 316.—Ic.: Nikitin, 
l.c. plate XLII. 

Annual. Plant (1.5)2—5 cm tall, with several or numerous scapes 
usually shorter than leaves, rarely almost as long. Leaves narrowly 
linear or lanceolate-linear, 2-4 mm broad, gradually narrowed toward 
base, more or less densely lanate beneath, glabrous or sparsely hairy 
above. Scape more or less densely tomentose, in the beginning erect, 
incurved in fruit. Spikes dense, capitate, 5-8 mm in diameter. Bracts 
with distinct herbaceous keel, broadly membranous along margin; lower 
bracts ovate or orbicular-ovate, glabrous, rarely hairy, upper narrower, 
often more or less hairy. Calyx lobes elliptic or ovate, 3.5-4.5 mm 
long, with distinct herbaceous keel, broadly membranous along margin, 
more or less densely lanate. Corolla lobes ovate or oblong-ovate, 1.5— 
2 mm long, glabrous; corolla tube densely villous. Capsule ellipsoidal, 
about 3 mm long. Seeds elliptic or oblong-elliptic, about 2.5 mm 
long, with 1 side convex and other deeply concave. April to June. 

On rubbly and clayey slopes, takyrs [clayey soils in deserts] saline 
soils.—Soviet Central Asia: Kyzyl-Kum, Kara-Kum, Amu-Darya. 
General distribution: Iran-Afghanistan (Afghanistan). Described from 
Kyzyl-Kum. Type in Leningrad. 


25. P. lagocephala Bge. in Mém. sav. étrang. Acad. Sc. Pétersb. 
VII (1851) 445; O. and B. Fedch. Perech. rast. Turkest. V, 196; Pilger 
in Pflanzenr. H. 102, 356; Nikitin, in Fl. Turkm. VI, 316.—Ic.: Nikitin, 
l.c. plate XLII. 

Annual. Plant 2(3)—10(12) cm tall, with several (2-10) obliquely 
erect, rarely erect or ascending scapes. Leaves linear-lanceolate, 
lanceolate or oblanceolate, (1.5)2—6(8) mm broad, gradually narrowed 
toward base, acute, more or less densely villous. Scapes more or less 
densely villous. Spikes dense, compact, ovate or cylindrical, 0.5-—2.5(3) 
cm long, densely villous. Bracts ovate or broadly ovate, with distinct 
herbaceous keel, margin broadly membranous, as also calyx lobes, 
densely villous. Calyx lobes oblong or ovate-oblong, about 3 mm 
long, with distinct herbaceous keel, margin membranous but not broad. 
Corolla lobes deltoid-ovate, about 1 mm long, densely villous (hairs 
longer than lobes), corolla tube glabrous. Capsule ellipsoidal, 3—3.5 


162 


154 


mm long. Seeds oblong-elliptic or oblong-ovate, 2.5—3.5 mm long, 
with one side convex, the other deeply grooved. April to May (Plate 
VII, fig. 2). 

On clayey and rubbly slopes, shingles, takyrs [clay soils in deserts], 
saline soils (in plain clayey deserts).—Soviet Central Asia: Aralo- 
Caspian (Ust-Urt), Kyzyl-Kum, Kara-Kum, Amu-Darya, Pamiro-Alai 
(in south). Endemic (probably occurs in Afghanistan). Described from 
northern Kyzyl-Kum. Type in Leningrad. 


Subgenus 2. Psyllium (Juss.) Harms in Pflanzenfam. IV 3b 
(1895) 373.—Psyllium Juss. Gen. pl. (1789) 90, pro gen.— Plant with 
well-developed foliate stems. Leaves opposite, often bearing shortened 
shoots in axils. Subgenus has one section. 


Section 12. PsyLLiuM (Juss.) Barneoud, Rech. Level. struct. gen. 
classif. Pflantag. Plumbag. (1844) 19.—Psyllium Juss. Gen. pl. (1789) 
90 gen.—Capsule 2-seeded. Seed with one side grooved. Corolla 
glabrous, tube often transversely rugose. Anterior calyx lobes different 
from the posterior. Spikes short and dense. Annual and perennial herbs 
or subshrubs. Pubescence often glandular. 


26. P. indica L. Syst. nat. ed. 10, II (1754) 896; Sp. pl. ed. 2, 167; 
Willd. Sp. pl. 1, 651, Pilger in Pflanzenr. H. 102, 418; Kryl. Fl. Zap. 
Sib. X, 2568.—P. psyllium L. Sp. pl. (1753) 115, non in Sp. pl. ed. 
2, nec in Syst. nat. ed. 10.—P. ramosa (Gilib.) Aschers. Fl. Prov. 
Brandenb. (1864) 547; Hayek in Hegi, Ill. Fl. VI, 1, 143; O. and B. 
Fedch. Perech. rast. Turkest. V, 197; Grossh. Fl. Kavk. IV, 16.—P. 
arenaria Waldst. et Kit. Descr. and Ic. Pl. rar Hung. I (1802) 51; M.B. 
Fl. taur.-cauc. I, 111; Ldb. Fl. Ross. III, 486; Schmalh. Fl. II, 352.— 
Ic.: Waldst. et Kit. l.c. tab. 51; Hegi. l.c. tab. 246; Pilger, l.c. f. 42.— 
Exs.: GRF, No. 131 (sub P. arenaria Waldst. and Kit.). 

Annual. Stem usually erect, (5)20-60 cm tall, more or less 
branched, with opposite, sometimes fairly long branches and, besides, 
with shortened shoots in axils of leaves, rarely simple, sparsely 
pubescent with simple, as well as short glandular hairs. Leaves narrowly 
linear, linear or lanceolate-linear 1-3(4) mm broad, subobtuse, sparsely 
pubescent with simple, and sometimes also with glandular hairs. Spikes 
dense, compact, oblong-ovate or oblong-elliptic, 0.5—1.5(2) cm long, 
on axillary 1-5(6) cm long peduncles, pubescence similar to stem. 
Two lower bracts differing sharply from the remaining in shape, broader 
in base, orbicular-ovate, narrowly membranous along margin, suddenly 
narrowed at apex into linear-subulate 3-10 mm long appendage, more 
or less densely pubescent, upper bracts more or less keeled, broadly 


163 


155 


obovate, or almost spatulate, obtuse, broadly membranous along margin, 
herbaceous keel more or less pubescent. Calyx lobes about 4 mm long, 
sparsely pubescent, fimbriate along margin, dissimilar, 2 anterior ones 
obovate or elliptic-ovate, obtuse, posterior 2 ovate-lanceolate, acute, 
more or less keeled. Corolla lobes ovate-lanceolate, acute. Capsule 
broadly elliptic, obtuse, 3—3.5 mm long, bilocular. Seeds oblong-elliptic, 
brownish-black, shiny, about 2.5 mm long, with one side convex, the 
other sulcate. June to August. 

On sands, sandy-clayey terraces, along roadsides, and in weedy 


places.— European USSR: Ladoga-Ilmen (rarely), Baltic Region, Upper 


Volga, Volga-Kama, Upper Dniester, Upper Dnieper, Middle Dnieper, 
Volga-Don, Trans- Volga, Bessarabia, Black Sea Region, Crimea, Lower 
Don, Lower Volga. Caucasus: All regions. Western Siberia: Upper 
Tobol (southern part), probably in southern Irtysh. Soviet Central Asia: 
Aralo-Caspian, Balkhash Region, Tien Shan (rarely). General 
distribution: Scandinavia (rarely, introduced), Central and Atlantic 
Europe, western and eastern Mediterranean, Balkans-Asia Minor, 
Armenia-Kurdistan. Described from Egypt. Type in London. 


27. P. psyllium L. Sp. pl. ed. 2 (1762) 167; Pers. Syn. I, 140; 
Boiss. Fl. or IV, 891; Grossh. fl. Kavk. IV, 16; Pilger in Pflanzenr. H. 
102, 422.—Ic.: Pilger, l.c. f. 43. 

Annual. Stem erect, rarely ascending, 5—20(40) cm tall, simple or 
more or less branched (sometimes with long ascending branches at 
base of stem), usually with shortened shoots in leaf axils, more or less 
pubescent with simple and glandular hairs. Leaves narrowly linear, 
linear or lanceolate-linear, 1-3(4) mm broad, subobtuse, somewhat 
expanded at base, sparsely pubescent. Spikes dense, compact, elliptic 
or ovate, 0.5-1(1.5) cm long, on axillary 1-4(5) cm long peduncles, 
more or less densely pubescent, stunted forms with a single terminal 
inflorescence; inflorescences usually several, often crowded at apex of 
stem and branches. Lower bracts ovate or ovate-lanceolate, membranous 
along margin, gradually narrowed at apex into oblong-linear fairly 
long herbaceous appendage, more or less pubescent; upper bracts 
resembling lower, often broader and with shorter appendage. Calyx 
lobes 3-4 mm long, more or less pubescent, dissimilar, 2 anterior ones 
oblanceolate, 2 posterior ovate-lanceolate and oblong-ovate, slightly 
acuminate. Corolla with fairly long, somewhat rugose tube and ovate, 
acuminate, about 2 mm long lobes. Capsule broadly elliptic, obtuse, 
3-3.5 mm long, 2-seeded. Seeds dark brown, oblong or elliptic-oblong, 


- 2.5 mm long, with one side convex, the other sulcate. April to May. 


On sands and dry and rubbly slopes, as well as weedy places.— 
Caucasus: Eastern Transcaucasia. Soviet Central Asia: mountainous 


164 


156 


Turkmenia. General distribution: Eastern and western Mediterranean, 
Balkans-Asia Minor, Armenia-Kurdistan, in parts of Iran; occasionally 
introduced in Central Europe, as well as in South America. Described 
from Southern Europe. Type in London. 

Economic importance. Used in treatment of gastric disorders. 


GENUS 1382. Littorella! Bergius 


In Vet. Acad. Handl. XXIX (1768) 341. 

Flowers unisexual, monoecious, forming several axillary 
inflorescences. Staminate flowers usually solitary (rarely in twos), 
with rudimentary pistil, commonly situated on long pedicel. Calyx 
lobes 4. Corolla with 4-lobed limb. Stamens 4, with long filaments. 
Pistillate flowers 2—8(10), in axils of bracts at base or in middle part 
of pedicel of the staminate flower. Calyx lobes 2-4. Corolla with 
short, dissimilar lobes. Pistil 1, with bilocular ovary. Fruit—indehiscent 
1-seeded nut. Perennial, littoral and aquatic herbaceous plants with 
rosette forming leaves. 

This genus contains three species: one in Europe, another in North 
America and a third in South America (A. Donat in Pflanzenr. 3 
Reihe, Heft 8). 


1. L. uniflora (L.) Aschers. Fl. Prov. Brandenb. (1864) 544; Pilger 
in Pflanzenr. H. 102, 433.—Plantago uniflora L. Sp. pl. I (1753) 
115.—L. lacustris L. Mant. Pl. If (1771) 295; Ldb. Fl. Ross. II, 475.— 
Ic.: Hegi, Ill. Fl. VI, 1, tab. 246; Pilger, I.c. f. 44.—Exs.: Pl. Finl. exs. 
Nos. 950-951. 

Perennial. Plant 2—12(15) cm tall, with shortened rootstock, 
creeping shoots (stolons) and narrow, linear or linear-subulate leaves, 
forming rosette. Terrestrial plants with leaves more numerous and 
shorter, dorsiventral, sulcate above, sometimes hairy. Staminate flowers 
on 2—7(9) cm long pedicel, bearing in the middle membranous, ovate 
bracts. Calyx lobes or staminate flower elliptic-lanceolate, 4—5(6) mm 
long. Corolla tube about 4 mm long; corolla lobes elliptic-ovate, about 
2 mm long. Pistillate flowers 2(3) in number. Bracts deltoid-lanceolate 
at base of pedicel of pistillate flower. Calyx lobes 2(3), resembling 
bracts, lanceolate, about 3 mm long. Corolla tubular, about 4 mm 
long. Pistil with long style. Ovary bilocular, but with only 1 ovule. 


'From Jittus—bank. All species of Littorella grow in water in the littoral zones of 
water reservoirs as well as along their banks. 


165 


157 


Nut cinnamon brown, elliptic, 2—2.5 mm long, tuberculate-rugose. May 
to September. 

In water of the littoral zone of reservoirs, at depth of 0.2—2(2.5) 
m, as also along damp banks. Submerged plants usually do not flower, 
although they develop flower buds. European USSR: Ladoga-Iimen, 
Baltic Region, extends to Upper Dnieper. General distribution: 
Scandinavia, Central and Atlantic Europe, western Mediterranean 
(rarely), the Azores. Described from Western Europe. Type in London. 

Note. In the experiments of Visloukh [Izv. Gl. bot. sada SSSR, t. 
29, vyp. 5-6 (1930)], under semiflooded conditions of cultivation, L. 
uniflora produced bisexual (instead of staminate) flowers at the end of 
May and in June, whereas it produced staminate flowers in July and 
August. The bisexual flowers in June were sterile. 


Order 38. RUBIALES ENGL. 


Flowers typically 5—4 merous, actinomorphic or zygomorphic with 
equal or unequal stamens. Ovary inferior, multilocular or unilocular. 
Ovules pendulous, one to many in each locule. Herbs, often woody at 
base and small shrubs with opposite, usually undivided leaves. 


Family CLI. RUBIACEAE’ JUSS. 


Flowers regular, almost always bisexual. Calyx adnate with ovary, 
its limb usually rudimentary, rarely in form of teeth of different shapes 
and size at apex of ovary and fruit. Corolla tubular-infundibuliform, 
infundibuliform, campanulate or rotate, its limb having 3-5(6) segments 
or lobes with or without appendage at apex. Stamens 3—5(6), alternating 
with segments or lobes of corolla. Ovary inferior, bilocular, rarely 4- 
locular (Mitchella L.), with 1, rarely (genus Karamyschewia Fisch. 
and Mey.) several ovules in each locule; style one, bipartite at apex, 
with 2 usually capitate stigmas, rarely (genus Phuopsis (Griseb.) Hook. 
f.) stigmas grow into a single clavate stigma, barely bipartite at apex. 


‘Treatment by I.A. Linczevsky (characteristics of family, subfamilies, and tribes, 
key to genera, and genera Karamyschewia Fisch. and Mey., Gaillonia A. Rich., Sherardia 
L., Crucianella L., Phuopsis (Griseb.) Hook. f., Valantia L.), E.G. Pobedimova (genus 
Asperula L., except sections Cynanchica, Sherardiana and Oppositifoliae, genera Galium 
L. and Microphysa Schrenk), M.V. Klokov (section Cynanchica of genus Asperula L.), 
A.I. Pojarkova (genus Rubia L.), B.K. Schischkin (genera Callipeltis Stev., Leptunis 
Stev., Mitchella L. and part of genus Asperula L.). 


158 


Fruit—binate, dry nut, with separable or inseparable halves (mericarps) 
when ripe or (genus Rubia L.) more or less succulent, drupaceous, 
sometimes (genus Valantia L., Callipeltis Stev.) simple (because of 
rudimentary second ovule of the ovary) dry nut or (genus 
Karamyschewia Fisch. and Mey.) bilocular capsule. Annual or perennial 
herbs, rarely undershrubs and shrubs with opposite branches, opposite 
or pseudoverticillate (because of development of large leaflike stipules) 
leaves and small flowers, sometimes solitary, often in lateral or terminal 
cymes, usually forming paniculate, spicate or capitate inflorescence. 

The family comprises about 400 genera and over 5,000 species, 
the overwhelming majority of which belong to the tropical flora of 
both hemispheres and are numerous, sometimes large trees and shrubs. 
Among the useful tropical plants of this family, mention should be 
made of coffee (genus Coffea L.) and cinchona (genus Cinchona L.); 
species of the genus Rubia L. (see p. 363) in the flora of the USSR, 
which provide stable dyes for natural fibers, have some importance as 
dye plants. 

The division of the family Rubiaceae into the subfamilies and 
tribes in the present treatment follows the scheme of Schumann [E.-Pr. 
Natiirliche Pflanzenfam. IV Teil, Abteil, 4, 5 (1897)]. Schumann’s 
scheme has recently been revised. Bremekamp [B. Bremekamp. 
Huitiéme congrés international de Botanique, Rapports et 
communications aux sections 2, 4, 5 and 6, Paris (1954) 113-114] 
proposed a new division of the family into six subfamilies on the 
basis of criteria that are different from those adopted as the basis for 
the Schumann system. However, in his brief article, Bremekamp 
indicated only an outline of his new system without the pertinent 
details, and its use in the “Flora of the USSR” is fraught with many 
difficulties. 

The following [taxa] are known in the fossil state: Galium palustre 
L. (fruits—Upper Quaternary Period at settlement of Endovsky on 
the Khopér River (P.A. Nikitin); G. verum L.? (fruits)—-Lower 
Pleistocene of Lower Volga (P.A. Nikitin). Galium sp. (fruits) — 
Pliocene at village of Krivoborye on Don (Voronezh Region, P.A. 
Nikitin). 


KEY TO GENERA 
1. Corolla throat and often lobes, inside with tuft of hairs. Ovary 4- 


TOCUIAR 0. ccsscnccessnccnsascunssecrtnesaatte aatere a tceareect 1384. Mitchella L. 
+ Corolla throat glabrous. Ovary bilocular .............. cc ceecesseeeeeeees Be 


166 


167 


v1 


n 


Se 


159 


Fruit—bilocular capsule. Ovules and seeds several in locules of 
ovary and capsule ............. 1383. Karamyschewia Fisch and Mey. 
Fruit—binate or simple dry coccus or more or less succulent, 
drupaceous. Ovules and seeds in locules of ovary and in halves of 


fruit ‘alwaySiSeparatedi:. Carats. ce ened. ss dees. deabRatsa even cs denne ees 3. 
Leaves opposite, connate at base into a sheath, without large leaflike 
Stipules:.2.0.. 2.0... ok a eee 1385. Gaillonia A. Rich. 


Leaves pseudoverticillate, with large, leaflike stipules, sometimes 
opposite (without stipules), but connate at base into a sheath ....4. 
Calyx limb distinct, 6-dentate ...............ceee 1386. Sherardia L. 
Calyxelimibmn of distinct age Sater cae. | Rontd. eeacPatea tame nnenecesesee 3 

Corolla tubular-infundibuliform, infundibuliform or campanulate 
with distinct tube, exceeding or equaling corolla limb, rarely 
SOmle WNalSMORter 12 ..citieser ee). Pee Ee ee se eR cece dk 6. 
Corolla rotate, without distinct tube ..............cessecesscesseeeseeeseeeenes 11. 
Style long, far exserted from corolla. Stigmas connate into one, 
clavate, scarcely bifid at apex ... 1388. Phuopsis (Griseb.) Hook. f. 
Style not exserted from corolla. Stigmas not connate, always 
distinctly Panter ets. Src. s Cees. scacsead. . Dee edw de. pee tee -edaaasad Ts 

Flowers with bracts and paired bracteoles, situated decussately i in 
pelationito bracts <...0s-te8 sees Sees 1387. Crucianella L. 
Flowers with bracts, rarely isolated (unpaired) bracteoles ......... 8. 

Fruit more or less succulent, drupaceous .............. 1394. Rubia L. 
HPGUTE=—— PY COCCUS .2-.....<0n-e2-cercceeteeaoaessnneennnesebepeeiete «AGRE me. 2) 

Mericarps included in vesicular calyx connate at apex, with coarsely 
granular suntace 2b. Cass oe eer 1391. Microphysa Schrenk. 
Mericarps not surrounded by accrescent CalyX ...........cccceeeees 10. 
Mericarps claviform, thickened and incurved at apex. Leaves 
filiform, in whorls of 10-16. Annual ........... 1390. Leptunis Stev. 
Mericarps globose, ovoid or oblong, not thickened at apex, 
sometimes slightly falcate. Leaves not filiform, in whorls of 2- 
10(14). Usually perennials, sometimes woody at base, rarely 


annals). £.03 oxtunitus ad. bluosda comiciicccs pees 1389. Asperula L. 
Pedicels in fruit concrescent in threes, elongating and bending 
downward in arches. ...........csscessesseeseeseeseeeeees 1393. Valantia L. 
Pedicels not: conerescent im! fruit ...........<.----<o0ecce-0¢--seneeoesocenenbors 12. 


Fruit simple (not binate). Flowers nutant, covered by large, 
longitudinally folded, membranous bracts ..............::csscceeseeeesees 
X Prcn,.. sUsspe el get 00, NE lhe scien scl, wes 1395. Callipeltis Stev. 
Fruit binate or rarely simple due to nondevelopment. Flowers not 
nutant; without; large, Dracts,.:..2...........nes0not-adebeoqachnernnaasenndaanted- 13. 
Fruit—dry coccus. Corolla 4-partite ................ 1392. Galium L. 


168 


160 


+ Fruit drupaceous, usually more or less succulent, rarely dry. 
Corolla 5=6=partite 2. ee 1394. Rubia L. 


SUBFAMILY 1. Cinchonoideae K. Schum. in Pflanzenfam. IV, 4 
(1891) 16.—Ovary with several ovules in each locule. 


Tribe 1. OLDENLANDIEAE K. Schum. lI.c. 16.—Ovules erect, 
bracts finely (often almost setaceous) divided. 


GENUS 1383. Karamyschewia! Fisch. et Mey. 


Fisch. et Mey. apud Hohenack. in Bull. Soc. Nat. Mosc. XI (1838) 
266.—Karamyschewia Hook. f. in Benth. and Hook. f. Gen. Pl. II 
(1873) 59. pro subgen. Oldenlandiae.—Karamyschewia K. Schum. 
l.c. 26 pro sect. Oldenlandiae.—Theyodis A. Rich. Tent. fi. Abyss. I 
(1847) 364. 

Flowers bisexual. Calyx limb distinct, 8(10)-toothed. Corolla 
tubular, with 4-lobed limb equaling tube. Stamens 4. Style bipartite, 
stigma flattened. Fruit bilocular, many-seeded, indehiscent capsule, 
crowned at apex by accrescent calyx teeth. Small annual plants with 
opposite leaves and axillary as well as terminal flowers, usually 
gathered in small glomerules. 

Monotypic genus. 

Note. As seen from the synonyms, Karamyschewia has been 
described twice as a genus but, in either case, if it were united with 
the genus Oldenlandia L. s. 1., it would belong either to a monotypic 
subgenus or a monotypic section. The basic differences of the genus 
Karamyschewia from the genus Oldenlandia L. s. s. are: an 8(10)- 
toothed calyx limb and an indehiscent capsule. The genus Oldenlandia 
in its broad circumscription, as adopted by, for example, Schumann 
(K. Schumann, l.c.), including several isolated mono- and oligotypic 
sections, must, in our opinion, should be subjected to a critical review, 
and among the genera that should be restored undoubtedly ought to be 
the genus Karamyschewia Fisch. et Mey. 


1. K. hedyotoides Fisch. et Mey. apud Hohenack. in Bull. Soc. 
Nat. Mosc. IX (1838) 267; Ldb. FI. Ross. II, 395; Grossh. Opred. rast. 
Kavk. 246.— Oldenlandia hedyotoides Hook. f. in Benth. and Hook. f. 


‘In honour of A.M. Karamyschev (1744-1791), Russian naturalist, author of “O 
neobkhodimosti razvitiya estestvennykh nauk v Rossii [On the Necessity of Developing 
the Natural Sciences in Russia (1769)], who, in 1761-1766, was a student of Carolus 
Linnaeus in Uppsala. 


169 


161 


Gen. pl. II (1873) 59; Boiss. Fl. or. III, 11; K. Schum. in Pflanzenfam. 
IV, 4, 26; Grossh. Fl. Kavk. IV, 20; Mandenova in Fl. Gruzii. VIII, 
2.—Theyodis octodon A. Rich. Tent. fl. Abyss. I (1847) 364.— Exs.: 
Herb. Fl. Cauc. Nos. 532, 532b; Balansa, Pl. or. 1866, No. 1444; 
Letourn. Pl. aegypt. No. 262. 

Annual. Small marshy plant. Stems sometimes single, often several 
or numerous, procumbent-ascending, 5—20 cm long, small and unevenly 
branched, slender, sparsely pubescent at base, glabrous above, 
indistinctly 4-sided, with short, (0.5)1—1.5(3.5) cm long, internodes. 
Leaves opposite, oblong-linear or lanceolate, acuminate, with cuneate 
base, 1-1.5(2.5) cm long, 2~3(5) mm broad, glabrous above, sparsely 
pubescent beneath along midrib, base connate into membranous sheath, 
each side with 2-4 fine, subulate, 0.5—-1.5 mm long teeth (stipules). 
Flowers subsessile, axillary and terminal, usually in few-flowered small 
glomerules. Ovary obovoid, 1-1.5 mm long. Calyx limb with 8(10) 
linear-lanceolate, 0.5-1 mm long fimbriate teeth, some teeth in the 
middle considerably shorter and with narrow bases. Corolla white, 
tubular, about 1.5 mm long, with short, about 0.75 mm long, broadly 
cylindrical tube and 4-lobed limb, lobes shortly ovate-lanceolate, about 
0.75 mm long and 0.5 mm broad. Capsule truncate-obovoid, about 1.5 
mm long and 1.5 mm in diameter, sparsely pubescent, with 8(10) 
fimbriate, 1-2 mm long teeth at apex. Flowering June to October. 
Fruiting July to October (Plate IX, fig. 1). 

In lowlands—in rice fields, marshy places and moist river banks.— 
Caucasus: Western Transcaucasia (southern part), Talysh. General 
distribution: Baikans-Asia Minor (eastern part), Iran-Afghanistan (Gilan 
Province), Nile Valley, Abyssinia, Central Africa. Described from 
Talysh. Type in Leningrad. 


SUBFAMILY 2. Coffeoideae K. Schum. in Pflanzenfam. IV, 4 
(1891) 16.— Ovary with one ovule in each locule. 


Tribe 1. ANTHOSPERMEAE Cham. and Schlechtd in Linnaea, 
I1I (1828) 309; K. Schum. |.c. 16.—Ovule attached at base of ovary. 


GENUS 1384. Mitchella’* L. 


L. Sp. pl. (1753) 111.— Chamaedaphne Mitchell [Diss. Brev. Princip. 
Bot. (1784) 27] in Acta Acad. Nat. Cur. VIII (1784) app. 222.— 
Disperma J.F. Geml. Syst. nat. IT (1791) 892. 


‘Treatment by B.K. Schischkin. 
After John Mitchell, a doctor and botanist in Virginia, who died in 1768. 


170 


162 


Flowers regular, almost always bisexual. Calyx adnate with ovary, 
clayx limb in form of 3-6 teeth, persistent at apex of fruit. Corolla 
infundibuliform with 3—6-lobed, distinct limb, throat bearded. Stamens 
inserted in corolla throat, exserted. Ovary 4-locular. Fruit—berry. 

This genus includes two species, which are found in North America 
and Eastern Asia. 


1. M. undulata Sieb. and Zucc. in Abh. Acad. Muench. IV, 3 
(1846) 175; Hara, Enum. spermathophytarum japonicarum. II, 22; D. 
Vorob’ev, Mat. k. flore Kuril’skikh o-vov. 70.—M. repens L. var. 
undulata Makino in Bot. Mag. Tokyo, XXIII (1909) 69. 

Perennial. Rootstock slender, long. Stem prostrate on soil, rooting 
at nodes, glabrous, 5-25 cm long. Leaves opposite on short (1-5 mm 
long) petioles, entire, broadly ovate, truncated or indistinctly cordate 
at base, narrowed towards apex, obtuse, glabrous, S—20 cm long and 
4-12 mm broad, with prominent midrib and weakly noticeable lateral 
veins above, lateral veins more prominent beneath. Flowers in leaf 
axils, predominantly terminal, in twos, rarely solitary, on short pedicels. 
Calyx tube enveloping lower part of corolla, calyx teeth 3-6, persistent. 
Corolla white (according to Maksimovich), infundibuliform, tube about 
10 mm long, lobes 4 oblong, obtuse, 3 mm long and 1—1.5 mm broad; 
throat of corolla as well as lobes densely hairy inside. Stamens 4, 
inserted in corolla throat and slightly exserted. Fruit red (from 
Maksimovich’s label), globose berry 6-8 mm in diameter, with 2-8 
seeds. July to August. 

In oak-broad-leaved and birch forests.— Soviet Far East: Sakhalin 
(Kuril Islands—collected by D.I. Vorobev in Iturup and Urup). 
General distribution: Japan, Korea. Described from Japan. Type in 
Florence. 


Tribe 2. SPERMACOCEAE Spreng. Anleit. ed. 2, II (1818) 598; 
K. Schum. l.c. 16.—Ovule attached to ovary wall. Stipules 
when distinct from leaves (narrower and shorter than leaves) extremely 
finely (sometimes almost wooly) dissected or not. 


GENUS 1385. Gaillonia’ A. Rich. 
A. Rich. Mém. fam. Rubiac. (1830 separatim) 73 in Mém. Soc. Hist. 


Nat. Par. V (1834) 153.—Jaubertia Guillem. in Ann. Sc. Nat. 2 sér. 
XVI (1841) 60. 


‘In honor of Frangois-Benjamin Gaillon, French botanist (1782-1839). 


163 


Flowers bisexual, calyx limb distinct, 4-5 toothed. Corolla tubular- 
infundibuliform, with 4—5-lobed limb, lobes without appendage. 
Stamens 4-5. Style bipartite, stigma flattened. Fruit—dry, binate 
coccus. Semishrubs with opposite, linear leaves, connate at base 
forming sheath together with stipules, with flowers sessile at nodes 
and branch apexes in upper half of stems, forming more or less remote 
cymes. 

About 15 species, distributed from northwestern and eastern Africa 
(Algeria, Nubia, Island of Socotra) to Transcaucasia, southern Central 
Asia, Afghanistan and Pakistan, are known in the genus. 


1. Stems and fruit glabrous. Corolla 15-25 mm long .................... 
cabal nS Mire eae AEE ey! 1. G. bucharica B. Fedtsch. and Desjat. 
+ Stems and fruit pubescent. Corolla not more than 10-12 mm long 
peda oh tae it lt tessa OR Ue A ke AO id iD 
2. Corolla 3.5-4.5 mm long. Fruit densely villous ...................... 
aati. hme pe spite Seiad «Pega a 8 et 4. G. asperuliformis Lincz. 


+ Corolla 6-12 mm long. Fruit puberulent ......0000. eee 3. 
3. Corolla 10-12 mm long. Internodes almost always considerably 
shortemthantleaves Qucw. ths een Be os 4 2. G. szovitsii DC. 


+ Corolla 6-8 mm long. Internodes almost always longer than leaves 
PRES «tena, Salt ceo ESE RR ve Mend Ye 3. G. bruguierei A. Rich. 


Section 1. Microstepuus (Jaub. and Sp.) Boiss. Fl. or III (1875) 
13.—Microstephus Jaub. and Sp. Ill. pl. or. I (1842-1843) 138; pro 
subgen.—Calyx limb small, 5-partite, persistent, not accrescent. Corolla 
5-partite. Flowers in more or less regular cymes. 


1. G. bucharica B. Fedtsch. and Desjat. in B. Fedch. Rastit. 
Turkest. (1915) 708; B. Krech. in Fl. Turkm. VI, 321. 

Perennial. Undershrub, 40-60 cm tall, usually with thick, woody 
root and low, multiheaded caudex. Stems numerous, terete, slender, 
glabrous and smooth, woody and whitish towards base, dark green 
above, with opposite branches at each node, with 3-6 cm long 
internodes, almost always distinctly longer than leaves. Leaves 
opposite, sessile, narrowly linear, 1-3(4) cm long, and about 1 mm 
broad, dark green, sparsely pubescent, connate at base into a short 
white-membranous sheath, usually with 2—4 opposite, subulate teeth 
(stipules). Flowers at nodes and apexes of branches in upper half of 
stems. Ovary puberulent, oblong-obovate, 2—2.5 mm long. Calyx limb 
with 4—5 long-lanceolate, about 0.5—1 mm:long hairy teeth. Corolla 
white-gray (or dirty pink ?), about 15—20(25) mm long, fairly densely 
pubescent, with long narrowly obcuneate tube and 5-lobed limb, lobes 


172 


164 


linear-lanceolate, about 4-5 mm long and 1 mm broad. Coccus glabrous, 
oblong-obovate, about 4-5 mm long and 1.5-—2 mm in diameter. 
Flowering June. Fruiting July (Plate IX, fig. 2.). 
On outcrops of gypsum clays and sandstones in low mountains.— 
Soviet Central Asia: Pamiro-Alai (Kugitang Range, vicinity of Baisun, 
Babatag and Karatau ranges in southern Tadzhikistan). Endemic. 
Described from southern fringes of Karatau Range. Type in Leningrad. 


2. G. szovitsii DC. Prodr. IV (1830) 574 (“Sowitzii’’); Jaub. and 
Sp. Ill. pl. or. I, 139; Ldb. Fl. Ross. II, 396; Boiss. Fl. or. Ill, 14; 
Lipsky. Fl. Kavk. 332; Grossh. Fl. Kavk. IV, 21 and Opred. rast. 
Kavk. 246.—Exs.: Herb. Fl. Cauc. No. 242. 

Perennial. Undershrub 15—25(40) cm tall with fairly thick, woody 
root and low, multiheaded caudex. Stems numerous, terete, slender, 
scabrous with dense, tiny, whitish papillae, woody and whitish towards 
base, dark green above, with opposite branches at middle and upper 
nodes, with short (0.5)1—2.5(4) cm long internodes, almost always 
distinctly shorter than leaves. Leaves opposite, sessile, narrowly linear 
or linear-subulate, (1)1.5—3(3.5) cm long and about 1—-1.5 mm broad, 
dark green, and scabrous like stem, at base connate into fairly short, 
leathery membranous sheath, and in lower part usually with paired, 
linear-subulate and fairly long teeth (stipules). Flowers at nodes and 
ends of branches in upper half of stem. Ovary densely covered with 
tiny, oblong papillae, about 2 mm long. Calyx limb with 4—5-deltoid- 
lanceolate, 0.2-0.5 mm long teeth, fimbriate along margin. Corolla 
pinkish, about 10-12 mm long, densely covered with fine papillae, 
with narrowly obcuneate tube and S-lobed limb, lobes deltoid- 
lanceolate, about 2.5 mm long, and 1 mm broad. Cocci scabrous with 
fine papillae, oblong-ovate, about 3 mm long and 1.5 mm in diameter. 
Flowering June to August. Fruiting July to August (Plate IX, fig. 3). 

On dry stony mountain slopes at altitudes of 700-900 m.— 
Caucasus: Southern Transcaucasia (southernmost part of Araks Valley). 
General distribution: Armenia-Kurdistan. Described from vicinity of 
Nakhichevan. Type in Geneva, isotypes in Leningrad. 


3. G. bruguierei A. Rich. in Mém. Soc. Hist. Nat. Par. V (1834) 
153 (“Bruguierii”); Jaub. and Sp. Ill. pl. or. I (1842-1843) 139, 144; 
Boiss. Fl. or. III, 14; V. Krech. in. Flor. Turkm. VI, 320.—G. 
richardiana Jaub. and Sr. 1.c. 142.—G. incana Jaub. and Sp. I.c. 143.— 
Ic.: Jaub. and Sp. I.c. tab. 77, 75 (“G. richardiana”), 76 (“G. incana”).— 
Exs: Aucher-Eloy, Herb. or. No. 4679; Kotschy, Pl. Pers. bor. 1846, 
No. 563 (“G. incana’’); Bornm. It. pers.-turc. 1892-1893, No. 3582. 


175 


165 


Perennial. Undershrub, 15-25 cm tall, with thick, woody root and 
low, multiheaded caudex. Stems numerous, terete, fairly thick, densely 
covered with short, white reflexed bristles, woody and whitish towards 
base, grayish or dirty green above, with opposite branches at middle 
and upper nodes, short (1)1.5—2(3) cm long internodes, almost always 
longer than leaves. Leaves opposite, sessile, stiff and acicular, linear- 
lanceolate to narrowly linear, subulate at apex, 0.5(1)—1.5(3) cm long, 
and 1—2(4) mm broad, like stem densely and short bristly, connate at 
base into a more or less short leathery-membranous sheath, and in 
lower part usually with paired, linear or deltoid-subulate and fairly 
long, stiff, acicular teeth (stipules). Flowers at nodes and at apexes of 
branches in upper half of stems. Ovary densely covered with short, 
white bristles, oblong, about 2 mm long. Calyx limb with 4—5, deltoid, 
0.2-0.5 mm long teeth fimbriate along margin. Corolla pinkish, about 
6-8 mm long, densely covered with short, white bristles, with narrowly 
obcuneate tube and 5-lobed limb, lobes shortly linear-lanceolate, about 
1.5—2 mm long, and 1 mm broad. Coccus densely and extremely short 
bristly, oblong or oblong-ovate, about 3—4 mm long, and 1.5 mm in 
diameter. Flowering May to July. Fruiting June to August. 

On dry stony slopes, on shingles and conglomerates in piedmonts 
and low hills, at altitudes of 500-1000 m.—Soviet Central Asia: 
mountainous Turkmenia (Kopet-Dag, Kyurendag, Lesser Balkhan 
Mountains), Pamiro-Alai (northern foothills of Alai Range in the region 
of Isfara-Kokand). General distribution: Iran. Described from Iran 
(between Teheran and Isfahan). Type in Paris. 

Note. Specimens from the Pamiro-Alai are slightly different from 
Turkmenian and Iranian specimens (because of somewhat shorter 
pubescence in all parts of the plant and an even shorter corolla). The 
latter are, however, not entirely homogenous, differing in several minor 
features. Because the type material of G. bruguierei and of the 
apparently very closely related G. olivieri H. Rich., also described 
from Iran, were not available to me, I was unable to clarify the 
taxonomy of this group in more detail. 


4. G. asperuliformis Lincz. sp. n. in Addenda XXII, 689. 

Perennial. Undershrub 20--40(60) cm tall, with more or less thick 
woody, root and tall multibranched caudex. Stems numerous, terete, 
slender, shortly and more or less densely pubescent, woody and whitish 
below, dark-green above, with opposite branches at nodes, having up 
to 5—7 cm long internodes, many of which longer than leaves. Leaves 
opposite, sessile, narrowly linear, 1-2(3) cm long, and 0.5—1 mm broad, 
dark green, pubescent, connate at base into short white membranous 
sheath, with 4-6 opposite, shortly subulate teeth (stipules). Flowers in 


176 


166 


many-flowered cymes, at ends of branches. Ovary densely villous with 
white hairs, oblong-ovate, 1.5-2 mm long. Calyx limb with 5 ovate- 
lanceolate, more sparsely pubescent teeth, about 1 mm long. Corolla 
dirty pink (?), 3.5-4.5 mm long, fairly densely pubescent, with short 
about 2.5 mm long obcuneate tube and 5-lobed limb; lobes ovate- 
lanceolate or lanceolate, acuminate, about 1.5—2 mm long, and 0.5-1 
mm broad. Coccus densely villous with white hairs, oblong-ovate, about 
3 mm long, 1 mm in diameter. Flowering June to August. Fruiting July 
to August. 

On stony mountain slopes at altitudes of 1500-3000 m.— Soviet 
Central Asia: Pamiro-Alai (western Pamirs). Vanch, Bartang and 
Pyandzh river valleys—from Kurgovat to Lyangar-Kisht. General 
distribution: ?Iran-Afghanistan (?Afghanistan), India-Himalayas 
(Gilgit). Described from western Pamirs (valley of Pyandzh River 
between Dashtak and Kurgovat). Type in Leningrad. 

Note. This species of the affinity of G. eriantha Jaub. and Sp. is 
well distinguished from it by a much (almost 2 times) shorter and 
narrower corolla, longer and narrower leaves, thinner and taller stems, 
as well as a general habit reminiscent of certain species of Asperula, 
as a result of which it is sometimes confused with A. oppositifolia Rg]. 
and Schmalh. A. asperuliformis is the first representative of this unique 
group (perhaps a separate section of the genus) discovered in Soviet 
Central Asia. In this group, we include, besides this one and G. eriantha 
Jaub. and Sp., the northern Afghan G. dubia Aitch. et Hemsl. and also, 
apparently, G. puberula Balf. f. and G. thymoides Balf. f. from Socotra. 


Tribe 3. GALIEAE Dumort. Anal. famil. (1829) 33; K. Schum. in 
Pflanzenfam. IV, 4 (1891) 16.—Ovules attached to wall of ovary. 
Stipules leaflike (same shape and size and indistinguishable from leaf). 


GENUS 1386. Sherardia! L. 


L. Sp. pl. (1753) 102. 

Flowers bisexual. Calyx limb distinct, 6-toothed. Corolla 
infundibuliform, with 4-lobed limb, lobes without appendage. Stamens 
4. Style bifid. Stigma capitate. Fruit-dry, binate coccus. Annual 
herbaceous plant with procumbent stems and whorled, spatulate, and 
lanceolate leaves with flowers in capitate inflorescence, surrounded by 
sheath of leaves connate at base. 


‘In honor of William Sherard, English botanist (1659-1728). 


167 


Plate IX. 

173 1. Karamyschewia hedyotoides Fisch. and Mey., general appearance of plant, 
capsule.— 2. Gaillonia bucharica B. Fedtsch. and Desjat., general appearance 
of plant, flowers with stipules.—3. G. szovitsii DC.—4. Crucianella schischkinii 
Lincz., general appearance of plant, part of inflorescence.—5S. C. filifolia Rgl. 

and Wirkl., general appearance of plant. 5(a) part of inflorescence. 


177 


168 
Monotypic genus. 


1. S. arvensis L. Sp. pl. (1753) 102; M.B. FI. taur.-cauc. I, 99; 
DC. Prodr. IV, 581; Ldb. Fl. Ross. II, 396; Boiss. Fl. or. III, 19; K. 
Schum. in Pflanzenfam. IV, 4, 148; Aschers. in Ber. deutsch. bot. 
Ges. XI, 29; Schmalh. Fl. II, 6; Lipsky, Fl. Kavk. 332; Fedch. and 
Fler. Fl. 903; B. Fedch. Rast. Turkest. 710; Hayek. in Hegi, Ill. Fl. VI, 
1, 197; Grossh. Fl. Kavk, IV, 21; V. Krech. in Fl. Yugo-Vost. VI, 254; 
Grossh. Opred. rast. Kavk. 246; Klok. in Vizn. roslin URSR, 307; 
Mandenova in FI. Gruzii, VIII, 3; V. Krech. in Fl. Turkm. VI, 321.— 
Ic.: Fedch. and Fler. l.c. fig. 888; Aschers. l.c. tab. III, f. 1-11; K. 
Schum. l.c. f. 47, A-B; Rchb. Ic. Fl. Germ. XVII, tab. 132 
(MCLXXXIII); Hayek, l.c. tab. 247, f. 1, Bonnier, Fl. compl. Fr. 
Suisse et Belg. V, tab. 262, f. 1326.—Exs.: Fl. Cauc. exs. No. 170; 
Billot, Fl. Gall. et Germ. exs. No. 568; Hayek, FI. stir. exs. No. 788; 
Fl. exs. austro-hung. No. 2235; Bornm. It. Anatol. 1899, No. 4528. 

Annual. Plant with procumbent-ascending, branched, 4-sided, 5— 
40 cm long stems with faces sharply scabrous. Leaves sharply cuneate, 
lower-spatulate, 4 in a whorl, upper-lanceolate, 6 in a whorl, all sharply 
scabrous along margin. Flowers 6-8 in capitate cymes, surrounded by 
sheath of 8-10 stiff leaves connate at base. Calyx limb with 6 appressed 
hairy, sometimes (var. maritima Gris.) comparatively weakly developed 
teeth. Corolla 4—5 mm long, lilac or pink-lilac, sometimes (f. albiflora 
R. Keller) white, with long tube and broad 4-lobed limb. Coccus about 
4 mm long, scabrous, crowned at apex with 6-toothed, sometimes 
fairly weakly (only in form of small tubercles) expressed persistent 
teeth of calyx limb. Flowering April to June. Fruiting May to July. 

On dry slopes in scrub, in forest edges, often as weed in fields, 
old fields and in gardens. In Caucasus up to middle mountain belt.— 
European USSR: Karelia-Lapland (southern part, Petrozavodsk Region), 
Baltic Region, Ladoga-Ilmen (?), Upper Volga, Volga-Kama, Upper 
Dnieper, Middle Dnieper, Volga-Don, Volga Region, Crimea, Lower 
Don, Lower Volga (?). Caucasus: All regions. Soviet Central Asia: 
Said to be introduced to Turkmenia, but not confirmed by herbarium 
collections. General distribution: Scandinavia (southern part), Central 
and Atlantic Europe, western and eastern Mediterranean, Balkans-Asia 
Minor, Armenia-Kurdistan, Iran, Canary islands, North America 
(adopted). Described from Northern Europe. Type in London. 


178 


169 
GENUS 1387. Crucianella! L. 


o Sp. Pl. (1753) 108. 

Flowers bisexual, paired to solitary, situated in axils of bracts and 
paired bracteoles, decussate in relation to bracts. Calyx limb not 
distinct. Corolla infundibuliform, with long tube and 4—5-partite limb; 
corolla lobes linear or linear-subulate with incurved appendages of 
varying lengths. Stamens 4-5. Style bifurcate, stigma globose. Fruit— 
dry, double nutlet, easily divisible into 2 halves (mericarps). Annual or 
perennial herbs, stems sometimes woody below, with whorled leaves 
and solitary or paired flowers forming spicate or spicate-capitate 
inflorescence. 

In the genus are about 35-40 species, distributed along the 
Mediterranean coast from Spain and Morocco to Soviet Central Asia. 


Se Aerial! Plait ee, I RE ei i 2. 
+ Perennial plant with more or less thick, woody roots and stems 
somewhat woody in lower palt ............cccssssssessessecseeesseeseeeneetecees 11. 
2. Flowers borne singly (only in one of the paired axils of bracts) and 
extremely rarely or rarely forming spicate inflorescence .......... 3. 
+ Flowers in twos (in each paired axil of bracts) in lax or loose or 
more or less dense spicate infloresCeNCe ............:scseesceeeesseeeneeeees 5. 


3. Corolla highly (2%-3 times) exceeding bracts ............csceeeseeseees 
Ps SE. REALONE, IS. OL 1. C. schischkinii Linc. 
+ Corolla noticeably (but not more than one and a half times) 
exceeding or equaling bracts, or considerably shorter ............... 4. 
4. Corolla one and a half times as short as bracts .............scsseee 
NE SIRES OLIN LI SoBe 2. C. bucharica B. Fedtsch. 
+ Corolla considerably exceeding or equaling bracts ................... 
SSN ERE BERT BROT 3. C. sabulosa Korov. and Krasch. 
5. Bracts distinctly connate at base (Up to Y—Y2) ou... .eeeseseeseeseenes 6. 
+  Bracts scarcely connate at base or fF€€ ...........cceceeessesseseeeeeenees R. 
6. Flowers in lax, interrupted spicate inflorescences. Bracts small, 
about 3 mm long, with narrow white membranous margin ...... 
HSE MEAN PRUE RRL RS 1 RR 2 4. C. filifolia Rg]. and Winkl. 
+ Flowers in dense, closely and narrowly spicate inflorescences. Bracts 
large, about 6-7 mm long, with broad (almost to midrib) white 


membranous marpin’:.{ 220 aes 10. C. latifolia L. 
7. -Corolla 2 times as long as bracts ............ 5. C. divaricata Korov. 
+ Corolla scarcely exceeding or equaling bracts, or shorter ......... 8. 


‘From the Latin crux, a cross; from the cruciform position of the bracts and 
bracteoles relative to one another. 


179 


170 


8.  Bracts with broad (almost to midrib) white membranous margin 
over their entire length 2.0.50 eee 9. C. angustifolia L. 
+ Bracts with narrow or broad (but then only in lower half) white 
membranous margin or with somewhat thickened, narrow white 


chondroid margin .2...ciie lt. hscRese cel: Geslocieseds bas 9. 
9.  Bracts with narrow white membranous margin, narrowly lanceolate, 
long, almost aristate-acuminate ......... 6. C. baldshuanica Krasch. 


+  Bracts with broad white membranous margin, only in lower half or 
with somewhat thickened narrow white chondroid margin, broader, 

ovate or oblong-lanceolate, with short, subulate cusp ............. 10. 

10. Bracts with somewhat thickened, narrow, white chondroid margin, 
distinctly narrowed at base ... 8. C. chlorostachys Fisch. and Mey. 

+  Bracts with broad (almost to midrib) white membranous margin in 
lower half, scarcely narrowed at base ...... EiemanineaskhghAs 

oh RE RD, SOE, BR INES. a 7. C. exasperata Fisch. and Mey. 

11. Flowers in more or less sparse, narrowly spicate inflorescences, 
with small, 2-4 mm long bracts, at base almost always separated 

and not overlapping one another ................. 11. C. gilanica Trin. 

+ Flowers in dense and considerably broader, spicate inflorescences, 
with large, 6-12 mm long, bracts, always overlapping one another 


1 Sons REN, Be fil Moe ane it aan aaltepet 12. 
12. Corolla greenish-yellow, 7-9 mm long,, with small limb, about 
2-5-3: MM ACTOSS | yids estas. Sn eae Mee 12. C. suaveolens C.A.M. 


+ Corolla bright-raspberry red or pink, sometimes white, (10)12- 
14(16) mm long, with large limb, up to 5-6 mm in across 
Aha 5 abies, Jel claunts ately ss nae le ynented racials 13. C. sintenisii Bornm. 


1. C. schischkinii Lincz. sp. n. in Addenda XXII, 687. 

Annual. Plant glaucescent-green, 15-30 cm tall. Stem slender, 
fairly densely, divaricately dichotomously branched from the base or 
in upper half, branches more or less distinctly 4-sided, unevenly 
scabrous along the ribs or smooth, with fairly long internodes. Leaves 
in whorls of 4, flat, acute, margin fimbriate and scabrous, due to 
extremely short, whitish, rigid, spines, young leaves diffusely scabrous 
on both surfaces; lowest leaves lanceolate, perishing early, the rest 
narrowly linear, 1-2.5(4.5) cm long, and 1—1.5 mm broad. Flowers in 
distinctly isolated, rarely spicate, termina inflorescences, borne singly 
(only in one of the paired axils of bracts). Bracts lanceolate or linear- 
lanceolate, gradually subulate-acuminate at apex, with white 
membranous margin, densely fimbriate with short and rigid spines, 
connate at base to some height. Bracteoles equaling bracts, somewhat 
narrower, otherwise similar. Corolla with pinkish, glabrous, extremely 
narrow tube and green-yellow (?), short (1-1.5 mm) limb; lobes 


180 


171 


fimbriately setose along margin or rarely only at apex, 7-9 mm long, 
242-3 times longer than bracts; corolla lobes with almost as long linear 
appendage. Mericarp (almost mature) oblong-linear, scarcely expanded, 
about 3-4 mm long, brownish-chestnut, speckled with whitish dots, 
with micro-granulate surface. pre May. Fruiting May to June. 
(Plate IX, fig. 4). 

On sands.—Soviet Central Asia: Pamiro-Alai (sands in lower 
reaches of Surkhan Darya, Kdfirnigan and Vakhsh rivers). Endemic ? 
(probably can be found in adjoining parts of Afghanistan). Described 
from Surkhan Darya Valley between the town of Shirabad and village 
of Kokaity (perhaps from the Khuadag-Tau Plateau?) Type in 
Leningrad. 

Note. This species is related to C. bucharica B. Fedtsch. and C. 
sabulosa Korov. et Krasch., but differs sharply from both of them in 
having much (almost twice) shorter and broader bracts, much shorter 
spinules along margin of leaves and bracts; from C. bucharica, 
moreover, it differs in having a corolla twice as long and from C. 
sabulosa in a considerably narrower corolla. Do the plants from the 
sands of Kashka-Kum, in the lower reaches of the Vakhsh River (not 
fully developed), perhaps belong to a separate form? 


2. C. bucharica B. Fedtsch. in Fedde, Repert. sp. n. X (1912) 
464; V. Krech. in Fl. Turkm. VI, 324.—Ic.: In Krech. l.c. plate XLIV, 
f. dextra. 

Annual. Plant light or glaucescent-green, 5-20 cm tall. Stem 
slender, divaricately dichotomously branched usually from the base, 
branches more or less distinctly 4-sided, unevenly scabrous along ribs, 
with short internodes friable at nodes. Leaves in whorls of 4, flat, 
acute and with spiniscent tip, margin and often also along vein beneath 
fimbriate and sharply scabrous due to fairly long, whitish, rigid curved 
spines, lowermost lanceolate, perishing early, the rest linear-lanceolate, 
1—1.5(2.5) cm long and 1.4—2(2.5) mm broad. Flowers in unevenly 
separated and usually indistinctly isolated, rarely spicate terminal 
inflorescence, borne singly (only in one of paired axils of bracts), 
sometimes (toward bottom) oppositely in branches. Bracts subulate- 
linear, like leaves firabriate with rigid spines along white membranous 
margin and along vein beneath, scarcely connate at base. Bracteoles 
somewhat narrower, slightly shorter than bracts or (upper) as long. 
Corolla whitish (or whitish-yellow ?), glabrous, (2)3—4(5) mm long, 
one and a half times as short as bracts; lobes weakly developed, 
considerably shorter than appendage. Mericarp linear or scarcely 
expanded at apex, 4-5 mm long, brownish or grayish-cinnamon, 


183 


by2 


speckled with whitish spots, with microgranulate surface. Flowering 
April to June. Fruiting May to June. 

On sands.— Soviet Central Asia: Kyzyl Kum (southern half), Amu- 
Darya. Endemic. Described from Sundukli sands (near Mt. Saman by 
Lake Dengiz-Kul). Type in Leningrad. 


3. C. sabulosa Korov. et Krasch. sp. n. in Addenda XXII, 688; V. 
Krech. in fl. Turkm. VI, 324 (descr. ross.).—Ic.: v. Krech. l.c. plate 
XLIV, f. sinistra.—Exs.: H.F.A.M. No. 247 (“C. bucharica B. 
Fedtsch.”). 

Annual. Plant light green, 10—25(50) cm tall. Stem slender, 
indistinctly 4-sided or terete, simple or weakly dichotomously branched 
and more or less accumbent, glabrous and smooth, scabrous only 
towards base at places, with fairly long internodes. Leaves in whorl of 
4, flat, acute, margin -fimbriate and scabrous due to short, whitish, 
rigid curved spines, lowermost lanceolate, perishing early, the rest 
linear-lanceolate or linear, 1.5—2.5(3.5) cm long and 1-1.5(2) mm 
broad. Flowers, in distinctly isolated, extremely rarely spicate, terminal 
inflorescence, borne singly (only in one of the paired axils of bracts). 
Bracts subulate-linear, like leaves, fimbriate with rigid spines along 
white membranous margin, scarcely connate at base. Bracteoles slightly 
narrower than bracts, otherwise similar. Corolla with pink, glabrous 
tube and greenish-yellow limb, 8-10 mm long, distinctly longer 
(sometimes one and a half times) than bracts, rarely as long; corolla 
lobes fimbriately setose along margin or only at apex, linear appendage 
nearly as long as lobe. Mericarp linear or scarcely expanded at apex, 
4—5 mm long, brownish or grayish-chestnut and speckled with whitish 
spots, with microgranulate surface. Flowering April to May. Fruiting 
May to June. 

On overgrown sands.—Soviet Central Asia: Kara-Kum, Amu- 
Darya. Endemic. Described from Kara-Kum sands (vicinity of Uch- 
Adzhi [railway] station). Type in Leningrad. 

Note. This species is well distinguished from its closest relative, 
C. bucharica B. Fedtsch., by its glabrous and smooth stems with longer 
internodes, larger flowers exceeding or equaling the bracts (not shorter 
than one-and-a-half times), more remote and clearly isolated 
inflorescences and other characters. 


4. C. filifolia Rgl. et Winkl. in Regel in Izv. Obshch. lyubit. 
estestovzn. antrop. i etnogr. XXXIV, 2 (1882) 42; O. and B. Fedch. 
Perech. rast. Turkest. III, 137; Malinowski in Bull. Soc. Bot. Gen. 2 
sér. II (1910) 15; Korov. in Sched. ad H.F.A.M. X, 17; V. Krech. in 
Fl. Turkm. VI, 323.— Exs.: H.F.A.M. No. 249a (var. parviflora Korov.), 


173 


Plate X. 


181 1. Crucianella divaricata Korov., general appearance of plant, part of 
inflorescence.— 2. C. baldshuanica Krasch., general appearance of plant, part of 
inflorescence.—3. C. exasperata Fisch. and Mey., general appearance of plant, 
part of inflorescence.—4. C. chlorostachys Fisch. and Mey., part of 
inflorescence.—5. C. suaveolens C.A.M. —6. C. sintenisii Bornm.—7. Phuopsis 
stylosa (Trin.) Hook. f., general appearance of plant, flower with 

bracts. 


184 


174 


No. 249b (var. grandiflora Korov.); Sint. It. transcasp.-pers. 1900- 
1901, No. 253a. 

Annual. Glaucescent-green plant, 15-50 cm tall. Stem slender, 
indistinctly 4-sided or terete, sometimes simple, often sparsely 
divaricately dichotomously branched, glabrous and smooth, scabrous 
only towards base. Leaves in whorls of 4, glabrous, with revolute 
scabrous margin, lower leaves linear-lanceolate or broadly linear, 
perishing early, the rest linear, acute, 2-5 mm long, and about 1-1.5 
mm broad. Flowers in lax spicate, terminal inflorescences sparsely 
flowered below. Bracts ovate or ovate-lanceolate, short-acuminate, 
extremely short fimbriate along white membranous margin, elsewhere 
glabrous, distinctly (up to 1/4-1/3) connate at base. Bracteoles 
narrower, linear-lanceolate, almost equaling bracts. Corolla pink (or 
yellowish-green ?), glabrous or finely scabrous, 6-10 mm long, 2-3 
times longer than bracts; lobes with as long linear appendage. Mericarp 
narrowly obovate, 2—-2.5 mm long, brownish, speckled with whitish 
spots, with parvi-granulate surface. Flowering April to May. Fruiting 
May to June (Plate IX, fig. 5). 

Plains and low piedmonts, on clayey and sandy soils.— Soviet 
Central Asia: Kyzyl-Kum, Kara-Kum, mountainous Turkmenia 
(Badkhyz), Amu-Darya, Syr-Darya. General distribution: Iran- 
Afghanistan (northern Afghanistan). Described from Syr-Darya valley 
(“Chardara, sandy hills”) and from Kyzyl-Kum desert (at Baibek well, 
hilly sands). Type and paratype in Leningrad. 

Note. Two varieties are observed, adapted, as indicated, to different 
ecological conditions: 1) var. parviflora Korov. [in Sched. and 
H.F.A.M. X (1926) 17 (No. 249a)] with short bracts (6-7 mm long) 
exceeding corolla two times, with narrowly cylindrical tube, growing 
on clayey sands, and 2) var. grandiflora Korov. l.c. (No. 249b), with 
the corolla (9-10 mm long) three times longer than the bracts, with 
the tube noticeably expanded above, growing on hilly sands. Because 
of insufficient herbarium material of C. filifolia var. parviflora, it is 
difficult to define its geographical range accurately. However, given 
that C. filifolia var. parviflora does not occur in the regions lying to 
the west of Amu-Darya—Kara-Kum sands and Badkhyz, as well as 
northern Afghanistan, as may be judged from the fairly abundant 
material [of C. filifolia], it can be assumed that here we are dealing 
with the phenomenon of not merely ecological but also geographical 
vicariance. Hence, it is possible that C. filifolia var. parviflora 
represents a distinct, narrow geographic race. In order to confirm this, 
more herbarium material and field observations will be needed. It would 
also be advisable to study C. filifolia var. grandiflora in more detail. 


185 


175 


5. C. divaricata Korov. in Bot. mat. Gerb. Bot. Sada, V. (1924) 
iS} 

Annual. Glaucescent-green plant, 20-50 cm tall. Stem slender, 
indistinctly 4-sided or terete, divaricately dichotomously branched, 
glabrous and smooth, scabrous only toward base. Leaves in whorls of 
4, glabrous with revolute scabrous margin, lower leaves broadly 
lanceolate or lanceolate, perishing early, the rest linear or narrowly 
linear, acute, 1.5—3.5 cm long and about 1—1.5(2.5) mm broad. Flowers 
in spicate, lax, terminal inflorescences. Bracts narrowly lanceolate, 
long, almost aristate-acuminate, keeled, with fairly long spines along 
whitish membranous margin, in remaining part shortly and sparsely 
spinulose or glabrous, free at base. Bracteoles more narrow, deltoid- 
subulate, usually almost 2 times as short as bracts. Corolla pink, rarely 
also finely scabrous or almost glabrous, 7-10 mm long, 2 times longer 
than bracts; lobes with as long linear appendage. Mericarp oblong- 
ovate, about 2 mm long, brownish-chestnut, speckled with whitish 
spots, surface micro-granulate. Flowering May to June. Fruiting June 
(Plate X, fig. 1). 

In low mountains and middle mountain zone, at altitudes of about 
800-1800 m.— Soviet Central Asia: Pamiro-Alai (southern Tadzhikistan 
and southern Uzbekistan). Endemic. Described from Hissar Range 
(between Regar and Karatag) and Kugitang Range. Type and paratype 
in Tashkent; isotype in Leningrad. 

Note. There is a variety known as var. echinata Krash. (in Tr. 
Bot. inst. AN SSSR, 1, 3 (1937) 283), distinguished by its dense and 
long white-hispid pubescence on both surfaces of the bracts and, 
sometimes, shortly and sparsely hispid leaves, described from Tian 
Pass and from Dagana-Chakob. 


6. C. baldshuanica Krasch. in Tr. Bot. inst. AN SSSR, ser. 1, 3 
(1937) 283. 

Annual. Glaucescent-green plant, 15-35 cm tall. Stem slender, 
more or less distinctly 4-sided, divaricately dichotomously branched, 
glabrous and smooth, scabrous only towards base. Leaves in whorls of 
4, glabrous, with revolute scabrous margin, lower leaves lanceolate or 
ovate-lanceolate, perishing early, the rest narrowly linear, acute, 2-4 
cm long and about 1—-1.5 mm broad. Flowers in spicate, lax, terminal 
inflorescences. Bracts narrowly lanceolate, long, almost aristate- 
acuminate, sharply keeled, with white membranous margin, all glabrous 
or sometimes scabrous along keel, free or slightly connate at base. 
Bracteoles narrower, linear, almost 2 times as short as bracts. Corolla 
pink (?), finely scabrous, 7-10 mm long, equaling bracts or somewhat 
shorter; lobes with linear as long appendage. Mericarp (immature) 


186 


176 


oblong-obovate, about 1.5 mm long, with microgranulate surface. 
Flowering May to June. Fruiting June to July (?) (Plate X, fig. 2). 

In low mountains, at altitudes of 1000-1700 m.—Soviet Central 
Asia: Pamiro-Alai (southern Tadzhikistan). Endemic. Described from 
Baldzhuan Region and from Mergen-Kutal Pass, east of Kulyab. Type 
and paratype in Leningrad. 

Note. It is necessary to mention the form with the bracts about 
half as long—var. brevibracteata Lincz. (southern Tadzhikistan, 
between villages of Garab-Dara and Sary-Chashma, south of Kulyab, 
24, VII, 1935, No. 712. I.A. Linczevsky); there is, it seems, no 
distinction from the type in the remaining features. 


7. C. exasperata Fisch. et Mey. in Ind. IV Sem. Horti Petrop. 
(1837) 8; Hohenack. in Bull. Soc. Nat. Mosc. XI, 268; Ldb. Fl. Ross. 
II, 403, Boss. FI. or. III, 22; Lipsky, Fl. Kavk. 332; Korov. in Sched. 
ad H.F.A.M. X (1926) 15, p.p.; Grossh. Fl. Kavk. IV, 22 and Opred. 
rast. Kavk. 246; V. Krech. in Fl. Turkm. VI, 322.—C. hispidula Fisch. 
and Mey. in Ind. I Sem. Horti Petrop. (1835) 25; non Decaisne.— 
Exs.: Pl. or. exs. No. 122; H.F.A.M. No. 248a (var. glabrescens Boiss.). 

Annual. Glaucescent-green plant, 10-30 cm tall. Stem slender, 
more or less distinctly 4-sided, sometimes simple, often divaricately 
dichotomously branched almost from base or only toward tip, glabrous 
and smooth, scabrous only at base. Leaves in whorls of 4-6 (lowest 
ones in twos), glabrous or sparsely scabrous, with revolute, sparsely 
spinulose margin, lowest leave ovate, obtuse, perishing early, the rest 
narrowly linear, acute, 1-1.5(2.5) cm long and 1-2 mm broad. Flowers 
gently divaricate from more or less declinate bracts, forming linear, 
fairly dense, terminal spicate inflorescences somewhat more lax at 
base. Bracts ovate- or oblong-lanceolate, gradually acuminate and with 
short subulate cusp at apex, sharply keeled, fimbriate with long rigid 
hairs along margin, hairs shorter on veins, surface finely scabrous, 
scarcely narrowed at base, free, broadly (almost up to vein) white- 
membranous in lower part. Bracteoles narrowly lanceolate, somewhat 
shorter than or equaling bracts, otherwise similar. Corolla with whitish 
tube and pale yellow or reddish limb, about 2.5—-3 mm long, slightly 
longer, equaling or somewhat shorter than bracts; lobes with as long 
linear appendage. Mericarp oblong-ovate, 2—2.5 mm long, brownish- 
chestnut colored and speckled with whitish spots, with microgranulate 
surface. Flowering June to July. Fruiting July to August (Plate X, 
fig. 3). 

On dry, stony mountain slopes, at altitudes up to 2100 m (Kugitang 
Range).— Caucasus: Southern Transcaucasia, Talysh. Soviet Central 
Asia: Pamiro-Alai (Hissar Range and Kugitang), Tien Shan (western 


187 


177 


Tien Shan). Karatau and Kuraminskii ranges). General distribution: 
Balkans-Asia Minor (eastern part), Armenia-Kurdistan. Described from 
Talysh. Type in Leningrad. 

Note. A distinct form, annotated in the herbarium by I.M. 
Krascheninnikov as var. karatavica Krasch. is prevalent in the Karatau 
Range; it differs from the type and from var. glabrescens Boiss. (for 
which it is sometimes mistaken) in having entirely glabrous and smooth 
bracts, bracteoles and inflorescence axis. The foregoing specimens, 
issued [as exsiccatae] in “Herbarium Florae Asiae Mediae” (No. 248a), 
belong mostly to this variety and partly also to the typical one. 


8. C. chlorostachys Fisch. et Mey. in Ind. I Sem. Horti Petrop. 
(1835) 25; Hohenack. in Bull. Soc. Nat. Mosc. XI, 267; Ldb. Fl. Ross. 
II, 403; Boiss. Fl. or. III, 23; Lipsky, Fl. Kavk. 332; Grossh. Fl. Kavk. 
IV, 22 and Opred. rast. Kavk. 246.—C. chlorantha Fisch. et Mey. in 
Ind. IV Sem. Horti Petrop. (1837) 8, sphalm. nomen.—C. exasperata 
var. marginata Korov. in Sched. ad H.F.A.M. X (1926) 17.—Exs.: 
H.F.A.M. No. 248b (“C. exasperata var. marginata Korov.”). 

Annual. Light or glaucescent-green plant, 5—20(30) cm tall. Stems 
slender, distinctly 4-sided, sometimes simple, often more or less 
declinately dichotomously branched nearly up to base or only toward 
top, finely scabrous almost over entire length or up to middle, 
predominantly along ridges. Leaves in whorls of 4 or in twos 
(uppermost and lowermost), from finely and fairly densely scabrous 
to almost glabrous, with revolute, extremely sparsely and shortly 
fimbriate-spinulose margin; lowest leaves oblong-ovate, obtuse, 
perishing early, the rest narrowly linear, acute, 1—-1.5(2.5) cm long 
and about 1—1.5 mm broad. Flowers distinctly divaricate from more or 
less declinate bracts, forming fairly dense, terminal linear spicate 
inflorescence somewhat more lax toward base. Bracts oblong-lanceolate 
or lanceolate, gradually acuminate and with short, subulate cusp at 
apex, sharply keeled, besides slightly thickened whitish chondroid, 
fimbriate-spinulose margin, glabrous and smooth, distinctly narrowed 
and free at base, free. Bracteoles 2 times narrower, narrowly lanceolate 
or almost linear, distinctly shorter than bracts, otherwise similar. Corolla 
with whitish tube and pale (or greenish)-yellow limb, about 2—2.5 mm 
long, much shorter (up to one and half to two times) than bracts; lobes 
with short, subulate appendage. Mericarp oblong-obovate, about 2 mm 
long, brownish-chestnut colored and speckled with whitish spots, with 
microgranulate surface. Flowering May to June. Fruiting June to July 
(Plate X, fig. 4). 

On dry, stony mountain slopes, sometimes on fallow land, at 
altitudes of 800-1500 m.—Caucasus: Eastern and southern 


188 


178 


Transcaucasia. Soviet Central Asia: Pamiro-Alai (northern piedmonts 
of Alai range), Tien Shan (Mogoltau Range). General distribution: 
Balkans-Asia Minor (eastern part), Armenia-Kurdistan, Iran. Described 
from Transcaucasia (vicinity or Kirovobad). Type in Leningrad. 


9. S. angustifolia L. Sp. pl. (1753) 108; DC. Prodr. IV, 586; Ldb. 
Fl. Ross. II, 402, Boiss. Fl. or. III, 22; Schmalh. FI. II, 7; Lipsky, FI. 
Kavk. 332; Fedch. and Fler. Fl. 904; Grossh. Fl. Kavk, IV, 22 and 
Opred. rast. Kavk. 246; Mandenova in Fl. Gruzii, VIII, 4.—Ic.: Rchb. 
Ic. Fl. Germ. XVII, tab. 125 (MCLXXVI); Fiori and Paol. Ic. FI. Ital. 
390; Bonnier, Fl. compl. Fr., Suisse et Belg. V, tab. 263, fl. 1329; 
Javorka és Csapody, Ic. fl. hung. tab. 480, f. 3447.—Exs.: FI. exs. 
austro-hung. No. 2975 (C. oxyloba Janka); f. Schultz and F. Wint. 
Herb. norm. No. 56; Durieu. Pl. select. hisp.-lusit. No. 312. 

Annual. Glaucescent-green plant, 10-50 cm tall. Stems usually 
erect, fairly slender, more or less distinctly 4-sided, simple or with 
short and usually appressed branches, usually extremely finely scabrous 
along ribs. Leaves all finely and sharply scabrous with revolute margin; 
lower leaves in whorls of 4, linear-lanceolate, the rest in whorls of 6, 
linear, acute, 1-2(4) cm long and about 1-2 mm broad. Flowers gently 
divaricate because of more or less declinate bracts in overlapping 4 
rows, forming dense terminal linear spicate inflorescences. Bracts 
lanceolate or narrowly lanceolate, very gradually narrowed, almost 
aristate-acuminate, sharply keeled, with broad (almost up to midrib) 
white membranous margin, more or less densely, crenulately scabrous 
along margin and keel, free at base. Bracteoles distinctly shorter and 
narrower than bracts, otherwise similar. Corolla with whitish tube and 
yellow limb, about 4-5 mm long, somewhat shorter than or equaling 
bracts; lobes with short, linear appendage. Mericarp oblong-obovate, 
about 2.5 mm long, brownish-chestnut colored and speckled with whitish 
spots, with microgranulate surface. Flowering (May) June to July. 
Fruiting July to August. 

On dry, stony slopes, in shrubby overgrowths, from foothills to 
middle mountain belt.—European USSR: Crimea (southern coast). 
Caucasus: Ciscaucasia, western Transcaucasia (northern part), eastern 
Transcaucasia, Talysh. General distribution: Western and eastern 
Mediterranean, Balkans-Asia Minor. Described from Montpellier. Type 
in London. 

Note. Judging from their lanceolate or narrowly lanceolate, 
extremely gradually acuminate bracts, our specimens (both the Crimean 
and Caucasian ones) of this species are close to the form described 
from Hungary under the name C. oxyloba Janka [in Magy. Tud. Akad. 


189 


179 


Kozl. XII (1874) 162], treated by certain authors, however, as only a 
variety.— var. oxyloba (Janka) Halacsy [Consp. fl. Graec. I (1901) 
743]. 


10. C. latifolia L. Sp. pl. (1753) 109; DC. Prodr. IV, 586; Ldb. 
Fl. Ross. II, 402; Boiss. Fl. or. II, 20; Schmalh. Fl. II, 7; Lipsky, FI. 
Kavk. 332; Fedch. and Fler. Fl. 904; Grossh. Fl. Kavk. IV, 21.—C. 
monspeliaca L. Sp. pl. (1753) 109.—Ic.: Fedch. and Fler. l.c. fig. 
889; Rchb. Ic. Fl. Germ. XVII, tab. 126 (MCLXXVII); Sibth. et Sm. 
Fl. graec. II, tab. 139; Fiori et Paol. Ic. Fl. Ital. 390; Bonnier, FI. 
compl. Fr., Suisse et Belg. V, tab. 263, f. 1328; K. Schum. in 
Pflanzenfam. IV, 4, f. 47, C-D; Javorka és Csapody, Ic. fl. hung, tab. 
4890, f. 3447a.—Exs.: Fl. exs. austro-hung. No. 2234; Dérfl. Herb. 
norm. No. 5272; Bornm. It. syr. 1897. Nos. 728, 1910, 11898. 

Annual. Glaucescent-green plant, 10-35 cm tall. Stems more or 
less straight, fairly slender, distinctly 4-sided, simple or more or less 
divaricately branched, densely finely scabrous to almost glabrous. 
Leaves all finely and sharply scabrous, with revolute margin, lower 
leaves in whorls of 4, obovate or oblong, up to 1.5 cm long and 6- 
8 mm broad, middle and upper leaves in whorls of 4—6, from lanceolate 
to linear, 1-2(3) cm long and (1)1.5—2.5(3.5) mm broad. Flowers in 
linear spicate, dense and compressed (with densely appressed bracts), 
narrow, almost cylindrical, terminal inflorescences. Bracts broadly 
lanceolate or lanceolate, upper ones gradually acuminate and with 
short cusp, obtusely (almost indistinctly) keeled, with broad (almost 
from the midrib) white membranous margin, densely and finely 
fimbriate along margin, connate up to middle at base. Bracteoles much 
narrower than bracts, linear, almost as long, as also fimbriate along 
keel. Corolla with whitish, at tip yellowish or pinkish tube and yellow 
limb, about 6-7 mm long, much longer than bracts (hidden by them 
only up to half); lobes with as long linear-subulate appendage. Mericarp 
oblong-obovate to almost linear, about 2.5-3 mm long, brownish 
chestnut colored and speckled with whitish spots, with microgranulate 
surface. Flowering May to July. Fruiting July to August. 

On dry, stony slopes.— European USSR: Crimea (southern <oast). 
Caucasus (?) (reported by Ledebour, but not confirmed by recent 
collections). General distribution: Western and eastern Mediterranean, 
Balkans- Asia Minor. Described from Island of Crete and from 
Montpellier. Type in London. 


11. C. gilanica Trin. in Mém. Acad. Sc. Pétersb. VI (1818) 493; 
DC. Prodr. IV, 587; Ldb. Fl. Ross. II, 403; Malinowski in Bull. Soc. 
Bot. Gen. 2 sér. II, 16; Grossh. Fl. Kavk. IV, 21 in Opred. rast. Kavk. 


190 


180 


246; Ehrendf. in Ann. Naturhist. Mus. Wien, LVI, 214.—? C. orientalis 
Willd. ex Schult. Mant. III (1827) 215.—C. glauca A. Rich. ex DC. 
l.c. 586 (cum var.) and in Mém. Soc. Hist. Nat. Par. V (1834) 131; 
Malinowski, I.c. 16; V. Krech. in Fl. Turkm. VI, 323.—C. glauca a. 
genuina Boiss. Fl. or. HI (1875) 24.—C. glauca B. laxiflora Boiss. 
l.c. 25.—C. glauca var. gilanica Ktze. in Tr. SPb. bot. sada, X (1887) 
195; O. and B. Fedch. Perech. rast. Turkest. III, 137.—C. glauca B. 
ghilanica Lipsky. Fl. Kavk. (1899) 333.—C. glauca ssp. elbursensis, 
ssp. pontica (? ssp. remola, ssp. transcaspica, ssp. transcaucasica 
Ehrendf. I.c. 214—219.—? C. colchica Manden. in Zam. po sist. i 
geogr. rast. Bot. inst. AN Gruz. SSR, 17 (1953) 64 and in Fl. Gruzii, 
VIII (1952) 4 (descr. georg.).—Iec.: Trin. l.c. tab. 12; Mand. I.c. (1952) 
tab. 358.—Exs.: GRF, No. 1775; Sint. It. transc.-pers. 1900-1901, 
No. 485; pl. or. exs. No. 219. 

Perennial. Green or glaucescent-green plant, 20-50 cm tall with 
more or less thick, woody root. Stems numerous, fairly slender, 
indistinctly 4-sided or terete, more or less densely, rigidly puberulent, 
and scabrous to glabrous and smooth, base more or less procumbent, 
erect above, weakly branched usually only in upper part, upper 
internodes longer and lower shorter. Leaves in whorls of 4—6(8), surface 
usually finely scabrous, margin revolute, from lanceolate (in shady 
forms), linear and narrowly linear to nearly subulate, acute, fairly 
stiff, (1)1.5-2.5(3.5) cm long and 1-2(3.5) mm broad. Flowers in 
more or less sparse, narrowly spicate, terminal inflorescences of varying 
length. Bracts 2-4 mm long, ovate, oblong-ovate, or nearly lanceolate, 
more or less short, acuminate, boat-shaped-convex and keeled, with 
narrow white-membranous finely fimbriate margin, surface glabrous 
and smooth or pubescent and more or less scabrous, at base nearly free 
to more or less highly (sometimes almost up to middle) connate, lower 
in inflorescence always distinct, not overlapping one another. Bracteoles 
more or less equaling bracts, somewhat narrower, otherwise similar. 
Corolla dirty pink or greenish-yellow, 5-7 mm long, with short, 
pubescent or glabrous tube and small limb, about 2—2.5 mm across, 
much longer (2-3 times) than bracts; lobes with more or less as long 
linear appendage. Mericarp oblong-ovate, about 2.5—3 mm long, 
brownish-chestnut colored and speckled with whitish spots, with 
microgranulate surface. Flowering May to July. Fruiting June to August. 

On stony slopes in lower and middle mountain belts.— Caucasus: 
Western, eastern and southern Transcaucasia, Talysh. Soviet Central 
Asia: mountainous Turkmenia, Pamiro-Alai. General distribution: 
Balkans-Asia Minor (extreme east), Armenia-Kurdistan, Iran. Described 
from northeastern Iran (Gilan province). Type in London? (in British 
Museum, in Pallas Herbarium?). 


191 


181 


Note. The species is treated here in the broad sense, corresponding 
to that of Boissier (Boiss. l.c.), but under the prior name C. gilanica 
Trin. (instead of C. glauca A. Rich.). Only one attempt has been made 
so far (except for Boissier’s description of two varieties) to divide this 
species (or species aggregate) into geographic races. It was Ehrendorfer 
(l.c.), who described within C. glauca A. Rich. (accepting it as a 
distinct species, along with C. gilanica Trin.), six subspecies, 
distinguished in most cases by minor and, in addition (as it is evident 
from more complete material than that cited by Ehrendorfer), unstable 
characters. Of these subspecies, the more distinct is C. glauca ssp. 
kotschyi Ehrendf., related to C. suaveolens C.A.M. [? = C. sosnowskyi 
Grossh. Opred. rast. Kavk. (1949) 246, nomen] from eastern Turkey, 
as well as, perhaps, C. glauca ssp. pontica Ehrendf. (? = C. colchica 
Manden. l.c.) from western Transcaucasia and eastern Turkey. The 
remaining subspecies are much less distinctive. Ehrendorfer apparently 
did not have the opportunity to study the type of C. glauca A. Rich 
(at least he does not mention it) and did not indicate which of the 
subspecies is typical, and so, unfortunately, even the nomenclature 
remains unexplained to date. 


12. C. suaveolens C.A.M. Verzeichn. Pfl. Cauc. (1831) 52; Ldb. 
Fl. Ross. II, 403; Boiss. Fl. or. III, 25; Lipsky, Fl. Kavk. 333; 
Malinowski in Bull. Soc. Bot. Gen. 2 sér. II, 15; Grossh. Fl. Kavk. IV, 
21 and Opred. rast. Kavk. 246; Ehrendf. in Ann. Naturhist. Mus. Wein. 
LVI (1948) 214.—Exs.: Aucher-Eloy, Herb. or. No. 4666 and No. 
4669 (C. glauca A. Rich); Bornm. It. pers. 1902, No. 7078. 

Perennial. Glaucescent-green or glaucescent plant, 25-30 cm tall, 
with more or less thick, woody root. Stems numerous, fairly slender, 
whitish, distinctly swollen at lower nodes, indistinctly 4-sided or terete, 
from more or less rigidly puberulent to glabrous, more or less 
procumbent towards base, mostly weakly branched only towards top, 
with fairly long internodes, with much shorter internodes toward base. 
Leaves in whorls of 6—8, lower leaves often recurved, all more or less 
densely, finely and sharply scabrous, with strongly revolute margin, 
linear, acute, fairly stiff, (1)1.5—2(2.5) cm long and 1—1.5(2) mm broad. 
Flowers in dense and short, spicate or capitate-spicate inflorescences, 
narrowly paniculate in upper part of stem. Bracts lanceolate, 6-8 mm 
long, long acuminate-aristate, boat-shaped and sharply keeled, with 
narrow rigidly fimbrillate white membranous margin, surface glabrous 
or finely scabrous, free or slightly connate at base, all overlapping 
one another. Bracteoles, more or less equaling bracts, somewhat 
narrower, otherwise similar. Corolla greenish-yellow, 7-9 mm long, 
with finely scabrous or glabrous tube and small limb, about 2.5-3 mm 


182 


across, distinctly longer (up to one and a half to 2 times). than bracts; 
lobes with as long; linear appendage. Mericarp oblong-ovate, about 2— 
3 mm long, dark brown and speckled with whitish spots, with 
microgranulate surface. Flowering May to July. Fruiting June to August 
(Plate X, fig. 5). 

Stony slopes and rocky places in forest in middle mountain belt.— 
Caucasus: Talysh. General distribution: Armenia-Kurdistan, Iran- 
Afghanistan (northwestern part). Described from Talysh. Type in 
Leningrad. 


13. C. sintenisii Bornm. in Mittheil. Thiiring. Bot. Vereins. N.F. 
XVIII (1903) 49; O. and B. Fedch. Perech. rast. Turkest. III, 137; V. 
Krech. in Fl. Turkm. VI, 326.—Exs.: Ed. H.B.P. No. 94; Sint. It. 
transc.-pers. 1900-1901, No. 1768 (typus!). 

Perennial. Glaucescent-green or glaucous-green plant (24)40—70 
cm tall, with thick woody root. Stems numerous, thick, whitish, distinctly 
swollen at lower nodes, indistinctly 4-sided or terete, from more or 
less densely, rigidly puberulent to glabrous, more or less procumbent 
towards base, weakly branched usually only towards very tip, with 
fairly long internodes. Leaves in whorls of 6—8, all more or less densely, 
finely and sharply scabrous, with revolute margin, linear, acute, fairly 
stiff, (1.5)2—3(5) cm long and 1.5—2(3) m broad. Flowers in dense and 
short, capitate-spicate, inflorescences, arranged almost in corymbose 
or paniculate-umbellate form in upper part of stem. Bracts narrowly 
lanceolate, 8-12 mm long, aristate-acuminate, boat-shaped, and fairly 
sharply keeled, with finely and rigidly fimbrillate white membranous 
margin, surface diffusely fine scabrous (almost glabrous), free or slightly 
connate at base, all overlapping one another. Bracteoles distinctly 
shorter and narrower than bracts, otherwise similar. Corolla bright- 
raspberry red or pink, rarely white, (10)12—14(16) mm long, with 
somewhat inflated at tip, finely scabrous or glabrous tube and large 
limb up to 5-6 mm across, much longer (about one and a half to 2 
times) than bracts; lobes with almost as long linear appendage. Mericarp 
oblong-obovate, about 2 mm long, dark brown and speckled with whitish 
spots, with microgranulate surface. Flowering May to July. Fruiting 
June to August (Plate X, fig. 6). 

On mountain steppe slopes at about 1000-1800 m.— Soviet Central 

192 Asia: mountainous Turkmenia (western Kopet-Dag from Nukhur to 
Songu-Dag). General distribution: Iran-Afghanistan (northern 
Khorasan). Described from Mt. Syunt in western Kopet-Dag. Type in 
Weimar (?), isotypes also in Leningrad. 

Note. In describing this species, Bornmiiller (Bornm. I.c.) noted 
its isolated position in the genus Crucianella, and later [in Mitteil. 
Thiring. Bot. Vereins. N.F. XXIX (1912) 35], in a critical review of 


183 


the revision of the species of Crucianella L. by E. Malinovsky [in 
Bull. Soc. Bot. Gen. 2, sér. II (1910) 9-16], established the separate 
monotypic section Roseae Bornm. for it. Recently, from the vicinity 
of the town of Bodzhnurd in northern Khorasan, a new species was 
described, namely, C. khorasanica Ehrendf. [in Ann. Naturhist. Mus. 
Wien, LVI (1948) 219], whose author considers it to be very close to 
C. sintenisii Bornm. but at the same time relating C. sintenisii to the 
C. glauca A. Rich. and C. suaveolens C.A.M. “group.” 


GENUS 1388. Phuopsis! (Griseb.) Hook. f. 


Hook. f. in Benth. and Hook. f. Gen. pl. II (1873) 151.—Phuopsis 
Griseb. Spicil. fl. rumel. II (1844) 167, pro sect. Asperulae.— 
Laxmannia S.G. Gmelin ex Trin. in Mém. Acad. Sc. Pétersb. VI (1818) 
492, pro syn.—Nemostylis Stev. in Bull. Soc. Nat. Mosc. XXIX, 3 
(1856) 361, non Nutt. (1837, “Nemastylis”) nec. Herb. (1840, 
“Nemostylis’’). 

Flowers bisexual. Calyx limb rudimentary. Corolla tubular- 
infundibuliform with 5-fid limb, lobes without appendages. Stamens 
5. Style long, filiform, by far exceeding corolla; stigmas united, clavate 
slightly bipartite. Fruit—dry, binate, coccus, easily splitting into 2 
halves (mericarps). Perennial, herbaceous plant with whorled leaves 
and bright pink flowers in terminal capitate inflorescence. 

Monotypic genus, occupying an intermediate position between the 
genera Crucianella L. and Asperula L. 


1. P. stylosa (Trin.) Hook. f. in Benth. and Hook. f. Gen. pl. II 
(1873) 151; Grossh. Fl. Kavk. IV, 22 and Opred. rast. Kavk. 246.— 
Crucianella stylosa Trin. in Mém. Acad. Sc. Pétersb. VI (1818) 490; 
DC. Prodr. IV, 587; Ldb. Fl. Ross. II, 403.—Asperula stylosa Boiss. 
Diagn. pl. or. sér. 1, 3 (1843) 31 and FI. or. III (1875) 27; Lipsky, FI. 
Kavk. 333.—A. ciliaris DC. Prodr. IV (1830) 582.—Laxmannia 
fasciculata S.G. Gmelin ex Trin. l.c. 492; pro syn.— Nemostylis ciliaris 
Stev. in Bull. Soc. Nat. Mosc. XXIX, 3 (1856) 361.—Ic.: Trin. l.c. 
tab. XI; Bot. Reg. XXIV, tab. 55; Baill. Hist. pl. VII, 261; Gartenfl. 
XXXVI, tab. 529; Kerner, Zhizn’ rast. (Russian Ed.) II, figs. on p. 
252 and 254.—Exs.: GRF, No. 566; Aucher-Eloy, Herb. or. No. 4672; 
Bornm. It. pers. 1902, No. 7079. 


‘From Phu—a specific epithet of one of the species belonging to the genus 
Valeriana L., and opsis, meaning external appearance. 


193 


184 


Perennial. Plant 20-60 cm tall, usually with fairly slender, woody 
root. Stems numerous, fairly thick, but soft, sharply 4-sided and sharply 
scabrous from more or less numerous, recurved spines, suberect at 
base, simple, rarely branched. Leaves in whorls of (6)8—10, sessile, 
with slightly revolute, sharply scabrous margin, lanceolate or narrowly 
lanceolate, acute, 1-2(2.5) cm long and (2)3—5 mm broad. Flowers in 
terminal, many-flowered, almost globose head, surrounded by numerous 
leaves, resembling an involucre. Bracts ovate-lanceolate, long- 
acuminate, long and soft fimbriate along margin, 8-12 cm long. 
Bracteoles one and a half to two times shorter and narrower, otherwise 
similar to bracts. Corolla bright pink, 12-15 mm long, with large limb, 
up to 5S—6 mm acrass, lobes oblong-ovate or lanceolate. Style one and 
half times as long as corolla. Mericarps about 1.5 mm long, oblong- 
obovate, slightly flattened and with deep, elongated groove on the 
inner side, brown-chestnut colored, with narrow, whitish bands. 
Flowering May to August. Fruiting June (?) to August (Plate X, fig. 7). 

On mountain slopes, in forest edges and glades, up to middle 
mountain belt.—Caucasus: Talysh. General distribution: Armenia- 
Kurdistan (eastern part), Iran-Afghanistan (northwestern part). 
Described from northwestern Iran (Gilan). Type in Leningrad. 

Note. Kerner (l.c. p. 252-253, 295, 311) describes the extremely 
interesting details of the pollination process in this species. Maturation 
of the pollen takes place while the corolla is not yet open (in the 
‘bud’). The anthers dehisce before blooming, and the pollen settles on 
thickened warty end of the style—the stigma. The style gradually 
expands, raising the stigma, covered by pollen, to the arched crown of 
the unopened corolla, and at the time is in a state of such high tension 
that during the opening of the corolla and the rapid unfolding of the 
style, the pollen is ejected like a cloud of dust, which is strewn over 
the insects that usually cause the sudden opening of the corolla. Here, 
we have a typical case of the presence of a projectile mechanism 
together with protandrous dichogamy. 

Economic importance. Of interest as a beautiful, long-flowering, 
decorative plant for open ground; often grown in gardens. 


GENUS 1389. Asperula’” L. 


L. Sp. Pl. (1753) 103. 
Flowers bisexual, paired, with bracts but no bracteoles or rarely 
solitary (unpaired) bracteoles. Calyx limb rudimentary, rarely developed, 
'Treatment by E.G. Pobedimova, except section Cynanchica, treated by M.V. 


Klokov, and sections Oppositifoliae and Gerardianae, treated by B.K. Schischkin. 
7From Latin asper—meaning scabrous; several species have scabrous leaves. 


194 


195 


185 


3-6 toothed. Corolla tubulaf-infundibuliform or campanulate with well- 
developed tube exceeding or equaling limb, rarely somewhat shorter, 
with 4(5)-partite limb, lobes without appendage or with weakly 
developed appendage. Stamens 4(5). Style bipartite, stigma capitate or 
clavate. Fruit—dry, binate coccus, glabrous or pubescent. Perennial 
or annual herbs, sometimes with stems woody below, whorled rarely 
opposite leaves and small flowers, in clustered or corymbose heads or 
cymes, forming paniculate inflorescence. 

The genus is known to include more than 200 species, distributed 
mainly in the Mediterranean region. 

Note. While treating Asperula L. for the “Flora of the USSR,” we 
were convinced of considerable heterogeneity in the genus. We could 
treat some of the species as separate monotypic genera, as for example, 
Phuopsis Benth., Microphysa Schrenk and Leptunis Stev., but it is 
possible that certain species of the present genus Asperula also could 
be regarded as distinct genera. A monographic treatment of the genus 
Asperula L. on a worldwide scale is greatly to be desired. 

Economic importance. A red pigment has been discovered in 
the roots of various species of the genus Asperula. It is possible that 
these species of Asperula L. could have some significance as dye 
plants. f 


eye He awesiop posite cet mia ae eee cea, <a ee Di 
+ Leaves in whorls Of 4—10(14) 0.0... cceceessssseseteeeeeesessereeneesovees 4. 
2. Fruit and corolla, as well as entire plant, glabrous ..............:.:ssssseseeees 

prnetin benervers 2), then ncbeaes treater wheats. & 73. A. laevis Schischk. 
+ Fruit, as well as corolla, more or less pubescent ..................00+ 3. 
3. Fruit with long, dense pubescence, setae almost equaling diameter 
of hemiicarp 6..f.ce. ania. 71. A. oppositifolia Rg]. and Schmalh. 
Fruit with short, appressed pubescence .... 72. A. popovii Schischk. 
Airimuialyaetipesit tie fe set San cane ch tesa hat calc Sane tne ES ie 5. 
Perennialoftem: woodyjatibase at: Ske nena ike ee. cs i. 
Mericarps, 2—2.5 mm across (under microscope) finely papillose, 
usually without longitudinal white bands ......... 74. A. arvensis L. 
+  Mericarps 1-1.2 mm across, smooth, with white long bands .... 6. 

Corolla tube up to 6 mm long, shorter, equaling or slightly longer 

farina eh see 75. A. setosa Jaub. and Sp. 
+ Corolla tube 8—14 mm long, usually several times longer than bracts 
is Deh omen « antes hs seit sreees, Maen easy 76. A. azurea Jaub. and Sp. 
GorollasS=merous) ee aes es aie. 8. 
+ Corolla 4-, rarely 3-merous ...........ccccesessecsseeeseessecesseessasssneeseneens 16. 


my P+ 


ay 


196 


186 


so + 


10. 


11. 


12: 


13: 


14. 


LS: 


16. 


Leaves in whorls of 8—14. Inflorescence racemose, consisting of 
flower clusters aggregated in a head ..0........e eee eeseeseeeeneeeeeeeeees 
nS RR SR See, 1. A. molluginoides (M.B.) Rchb. 


Leaves: in whorlsiof 427 (2.4254. 2 a Ree Sees 9. 
Inflorescence spicate-paniculate, clusters of flowers opposite, 
distinct. Leaves green, with slightly involute margin .............. 10. 


Inflorescence spicate, interrupted, rarely almost uninterrupted. 
Leaves fleshy, grayish, involute along margin up to midrib .... 11. 
Stem tall, 40-50 cm. Leaves almost noninvolute along margin, 
elliptic, short, lax flower clusters (Great Caucasian range) ........ 
LEA RE EE Ne Ae atten 20 Ae Bs 3. A. albovii Mand. 
Stem shorter, 10-40 cm. Leaves weakly involute along margin, 
linear. Flower clusters more dense (Small Caucasian range and 


adjoining hills) .......... eles 2. A. prostrata (Adams) C. Koch. 
Corolla: glabrous outsides s2265 22.85. Ma a 12. 
Corolla densely and shortly pubescent or setulose .................. 13. 


Flowers grayish, very small. Leaves (4)7-9(10) mm long, 0.5—1 
mm broad. Bracts half or 1/3 as long as flowers. (Caucasus) ....... 
PRACT RSIS pee ates be Ae ATy 4. A. glomerata (M.B.) Griseb. 
Flowers purplish, larger. Leaves (5)10—12(18) mm long, (0.5)1(1.5) 
mm broad. Bracts 1/3 longer than flowers (Fergana, Pamiro-Alai) 
PRES OE Sh TS LIN Ee PONS Lae 6. A. ferganica Pobed. 
Corolla and stem setulose. Bracts broadly elliptical or broadly ovate, 
almost equaling flowers, without cilia at base ................seeeeeeeee 
Hees cdl abe siiawa ls DRUMS URE ais Sasi as TN 5. A. hirsutiuscula Pobed. 
Corolla and stem densely patently pubescent. Bracts lanceolate or 
ovate-lanceolate, long-ciliate, especially in lower part ............ 14. 
Flowers yeliowish, with cluster of long rigid cilia on bend of lobe 
tip. Bracts with extremely long, white, soft cilia along margin, 
longer than breadth of bracts ................8 7. A. pamirica Pobed. 
Flowers purplish, without cluster of rigid cilia on bend of lobe tip. 
Bracts, with short, thick bands along margin in or with long rigid 
erect hairs {neni ORR Ga ee AS. ST, 15. 
Stem 1.5—7 cm tall, plants pulvinate. Bracts several times shorter 
than flowers. Flowers terminal, on stems and branches, few in 
clusters. Leaves very small, 2-3 mm long, 0.5—0.75 mm broad 
8. ee. ee GRO RE AG ns eR 8. A. balchanica Bob. 
Stem 10-35 cm tall. Bracts somewhat shorter than or almost 
equaling flowers. Inflorescence spicate. Leaves larger, (4)7—8(11) 
mm long, (0.25)-0.5(1) mm broad ...... 9. A. turcomanica Pobed. 
Flowers with long and slender corolla tube, 3-4 times longer than 
lobes. Inflorescences monocephalous, terminal on stems and 


197 


17. 


18. 


£9: 


20. 


ou. 


24. 


187 


branches, surrounded by involucre. Leaves ovate or broadly ovate. 
(Section Monocephala Pobed.) ................:sssssessseesesseeseeeeeessees Poe Wie 
Flowers with shorter and broader corolla tube, equaling lobes or 
shorter, rarely 2-3 times as long, but then tube broad, whereas 


corolladobeswshort! e2cecti 2h, BRR RR 18. 
Anthers white, light brown when dry.....11. A. propinqua Pobed. 
Anthers dark brown when dry ................. 12. A. caucasica Pobed. 


Stems spreading or ascending. Branches almost from the very base 
of stem long, almost horizontal. Cymes few-flowered, axillary, 
Several Onl branches 2 etka i GRRE. 19. 
Stems erect, if weak and spreading, sometimes clinging or glabrous, 
but then plants small, with simple, about 3—4-flowered terminal 
cymes. Branches obliquely erect. Inflorescences terminal or in axils 
GbNpperMeavesionl yon OA, LA BRO, BR ies 20. 
Stems spreading, especially in lower internodes, densely patently 
white-pubescent, rarely scabrous, or glabrous. Cauline leaves (5)10— 
15(23) mm long, (1)2.5—3(5) broad, crowded on branches. Corolla 
tube shorter than lobes ..................... 69. A. humifusa (M.B.) Bess. 
Stems ascending, glabrous, lustrous of scabrous, rarely diffusely 
pubescent with white hairs. Cauline leaves large, (10)25-30(40) 
mm long, (1)3—4(5) mm broad. Corolla tube longer than or equaling 
TOBE SHOE LOSS. 70. A. insuavis Pobed. 


SCRE, EIGER, UTE IE BO CRE SY EG BAe 21. 

Cauline leaves in whorls of 4-10, if all whorls of 4, then ovate, 
broad, large! een, We ii BARA eS 22: 
Cauline leaves narrowly linear, mostly acuminate. Inflorescence of 
varied shapes. Fruits rugose, covered by hemispherical or cuneate 
tubercles, rarely flat, scaly outgrowths. (Section Cynanchina V. 
IRRECZ*) A EE IOS. ES RO UE, else asd naa atauascrecees 36. 
Cauline leaves elliptic, S-8 mm long, 2.5-3 mm broad. Cymes 
terminal, simple, 3—4-flowered, small. Alpine plants 5—15 cm tall 
ei) AMC ONS Td CMR Za 78. A. gracilis C.A.M. 
Inflorescences terminal, with 3 simple or divaricately or ternately 
EP ATIGHE CRETE 28 E1122). Mbs ater en ec eed DN ec tavcasemabestineearnwendee 23. 
Inflorescences: paniculate:s2 2 eek Se tees 24. 
Cauline leaves in whorls of 6—10, broadly oblanceolate, (18)40(54) 
mm long, (4)10(13) mm broad. Fruit with shiny hooked hairs, borne 
enstubercles 28. ndn tile, 12. O. odorata L. 

Cauline leaves in whorls of 6, basal leaves in fours, narrowly linear, 
(10)17—18(24) mm long, (0.5)1(1.5) mm broad. Carpels glabrous, 
with granulated surface .............ccseeeeseeees 77. A. taurica Paczoski. 
Cauline leaves in whorls of 6-8. Corolla lobes with short thick 
ARE NSA GP oa cn ans Nese cena eae cece acces cv ncxecestsanbvnn cwotaucmaaceoearseness 25. 


198 


188 


Zils 


28. 


29. 


34. 


Cauline leaves in whorls of 4—6. Corolla lobes obtuse or rounded 


Ab APO Kiy vansay snr suseessaagpastanel <ceak- Seep eee. deena eee oder deans, a2. 
Stems climbing, retrorsely spinulose along ribs ...................6+ 26. 
Sterns glabrous or pubescent with long, white hairs ................... 28. 


Flowers campanulate or tubular-infundibuliform, with broad and 
short tube, equaling lobes or somewhat longer. Staminal filaments 
2-3 times as long as elliptic or almost orbicular anthers .......... 27. 
Flowers tubular-infundibuliform, with short, narrow tube, 2 times 
as long as lobes. Staminal filaments 2 times as short as oblong 
AMtherS)assnsn-Saseont Supstere hee dean Ae Tees, steals 67. A. aparine M.B. 
Stems weak, spreading. Flowers snow white. Pedicels glabrous, 
equaling flowers or somewhat longer ..............ccsssccesseeesereeees 
uns. Loethaneas, ane cemomeye Rael, . rerees ur lig 66. A. rivalis Sibth. and Sm. 
Stems strong, erect. Flowers bluish or pale violet (darker in dry 
state). Pedicels retrorsely scabrous, shorter than flowers, equaling 
OVanyeecieesS ance ao, ates. a 68. A. karataviensis Pavl. 
Cauline leaves in whorls of 8-12, extremely narrow, filiform, 
(20)25(45) mm long, (0.25)0.5(0.75) mm broad. Inflorescence 
paniculate-corymbose, 1aXx ..........:csssseeees 56. A. octonaria Klok. 
Cauline leaves in whorls of 6—8, broader, (7)35(40) mm long, 
(0.25)1(1.5) mm broad. Inflorescence corymbose, narrower and 


SSS red Re INT, ass oewtencdscasonsineoneanan enue Mesa tyns seen euey anaes eemenaee tis 29. 
Cauline leaves long, (20)35(40) mm long, 0.5—1 mm broad, almost 
glabrous or scabrous along margin, due to fine setae ............ 30. 


Leaves shorter and broader, (7)15(25) mm long and (0.25)1(1.5) 
mm broad, strongly scabrous over entire surface due to fine setae 


Stemi clabrousenk.wk. Seema 2s 57. A. campanulata (Vill.) Klok. 
Stem densely pubescent on lower internodes and nodes .......... 
He ca steel Senet al donde anid tem 58. A. tyraica Bess. 
Stemyplabrous.-4..2..2ols. pk. BERG 59. A. galioides M.B. 
Cauline and lower leaves densely pubescent on lower internodes 
Daal erppy MeN a es ee 60. A. xerotica Klok. 
Cauline leaves with ring of short dense hairs at base ...............-- 
cosy aif Wwaceaamvene  Rtheebhoaes eereeeiate tees tae 65. A. maximowiczii Kom. 
Cauline leaves without ring of hairs at base ............:eseeeeee 33. 
Cauline leaves in whorls of 4, lower orbicular-ovate, rest ovate ... 
Macunads Spee Besevel cunenal a nck bene bined Ryne ee 64. A. platygalium Maxim. 
Cauline leaves in whorls of 6, lanceolate, ovate-lanceolate or linear 
sebyacirncet penetoet ae Reary oul Lepage sce. Nas. pligetine. senna es aieeR EAN a toe 34. 
Upper cauline leaves in whorls of 4, rest in whorls of 6. Corolla 3- 
partite, rarely 4-partite. Stemens 3, rarely 4 ... 63. A. tinctoria L. 


3S: 


36. 


ae 


38. 


41. 


189 


Lower cauline leaves in whorls of 4, rest in whorls of 6. Corolla 4- 
AMIR NS PARMETES AU ees, Refeckeeerateecuatens senatente cues «craves zetewaensnsncecs-nas=t 35), 
Corolla 1.3-2 mm long, panicle 8—25(35)-flowered. Stems 10—20 
cm tall with 4-5 whorls of leaves. Leaves usually declinate ........ 
< EERO ee PORTE, NBs BANE, oho 62. A. kryloviana Serg. 
Corolla about 3 mm long. Panicle many-flowered. Stems 20—50 
cm tall, leaves in whorls of 7-8, mostly obliquely erect ............ 
Bice curtilic Pa MPL e eae Cree ee EEE Eee ROPE eee 61. A. paniculata Bge. 
Fruit 1.5—2.5 mm, rarely up to 3 mm long, covered with extremely 
fine, hemispherical or cuneate tubercles. Plant chiefly characteristics 
of rubbly denudations, mostly in mountains, rarely in steppe ... 37. 
Fruit up to 3-4.5 mm long, covered by flat, scaly outgrowths, rarely 
almost smooth. Growing exclusively on sandy soil in plain forest 
steppe, Steppe Or SEMiIarid FEGIONS .............ceseeeceseseereereeseeteeeees TP, 
Corolla lobes almost as long as tube (a bit shorter or sometimes 
slightly longer). Corolla not longer than 4.5 mm ..................+- 38. 
Corolla lobes at most one and half times as short as tube. Corolla 
often longer than 4.5 mm (up to 8 MM) ...........ceeeeeeeeeeeeeee 46. 

Floral leaves linear or oblong-lanceolate, not broader than 1 mm, 
margin smooth. Flowers with distinct pedicels not forming dense 
clusters. Corolla whitish. Plants in southern (besides Crimea) and 
southeastern regions of European part of USSR ...............06 39. 
Floral leaves mostly ovate-lanceolate, broader than 1 mm, more or 
less serrate in basal part. Flowers almost sessile, terminal on few 
branches of inflorescence, in fairly dense 3—9-flowered clusters. 
Corolla distinctly pink. Exclusively found in mountain regions of 


MGR aes oA ite cxercscaosteansactenmene toes cer cantesbatmhet wes tbe cSt ane aes 44. 
Corolla scabrous-setulose. Pistil 1.5—2.25(2.5) mm long ........ 40. 
Corolla glabrous and smooth or nearly smooth (lobes sometimes 
diffusely setose along margin). Pistil 2-3 mm long ............... 43. 


Middle cauline leaves narrowly linear, flat, 0.75-2 mm_ broad. 
Almost entirely herbaceous plant, not forming mats, with flowering, 
stems up to 50-60 cm tall, characteristic of meadow steppe .... 41. 
Middle cauline leaves linear-setaceous, with strongly revolute 
margin, 0.25—-0.75 mm broad. Typically subshrub, forming lax or 
denser and pulvinate mats with flowering stems up to 20-30 cm 
tall, characteristic, exclusively, or calcareous denudations ..... 42. 
Stem mildly scabrous-setulose or scabrous in lower part. Corolla 
34.5 mm long. Pistil 2—2.5 mm long .......... 14. A. cynanchica L. 
Stem, to a considerable extent, densely pubescent with short but 
distinct setaceous hairs in lower part (together with leaves). Corolla 
2.25-2.75 mm long. Pistil 1.5—2 mm long .............cseeeeeeneeeeees 
mca uenneeannnnenntatnenete cod tnaa naka tenes tiassahcenuRh 13. A. semiamicta Klok. 


200 


190 


43. 


44. 


45. 


46. 


Subshrub, 3—20 cm tall, forming dense, often pulvinate mats. 
Flowering stems, scabrous-setulose throughout, fairly weakly 
branched only in upper part. Corolla 3—3.75 mm long, with sharply 
reflexed lobes. Fruit about 2 mm long ... 16. A. tephrocarpa Czern. 
Subshrub, 15-30 cm tall, forming lax mats. Flowering stems 
glabrous and smooth in upper half, branched from middle and 
below. Corolla 2.5-3 mm long, with less bent lobes. Fruit also 
somewinat samallen on. 252. s2isteocctendastitnsnaten 15. A. creticola Klok. 
Flowering stems, sharply scabrous throughout. Cauline leaves 
distinctly scabrous along margin. Dingy dark green, rigid plant 
with thicker stems, seen only in calcareous denudations along Volga 
F-YaTc i <4 50) 0) ee nn a ae 17. A. exasperata V. Krecz. 
Flowering stems mildly scabrous except for base, completely smooth 
at top. Leaves margin smooth. Bright green, lustrous, nonrigid 
plant, with thin stems, growing on exposures of various rocks, but 
not on calcareous rocks, in southern Cis- and Trans-Urals ........... 
“bos scutes aaueeetapawdhie oanesemmasate Bian ot Gates sores steer 18. A. petraea V. Krecz. 
Corolla small, 2.5-3 mm long, with about 0.5 mm broad 
lobes. Plant growing on calcareous denudations (known only 
from environs of Belogorsk) ...............:sesesseeeeees 19. A. supina M. 
Corolla 3.5-5 mm long, with 0.75-1.25 mm broad lobes. Plant 
growing in yaila (mountain pastures in Crimea) or southern 
MOUMEAIN SLOPES) lana cxnech saacdirerar sn bocidentney «ctl axceoeys saporcuambadarat eat 45. 
Flowering stems 5—15 cm tall, pubescent throughout atleast below 
nodes. Leaves up to 11 mm long, all scabrous-densely setose or, 
pubescent as in stem. Plant growing im yalia ................cscssesseeseeeenees 
senate ca asletleatlwsaaleN chad sick dotnet seu can cea eee ace 20. A. caespitans Juz. 
Flowering stems 10—20 cm tall, glabrous and smooth throughout or 
mildly scabrous only at base. Leaves up to 14 mm long, lower and 
middle leaves mildly scabrous-setulose along margin, only upper 
ones smooth. Plant growing in southern mountain slopes .... 
vocals oobtiaas encepagneaacmupppnas ts duazyatieas tec nepeamns 21. A. tranzschelii Klok. 


PUSH acvinacn.sakesraececasrecyenss tapos oneal cece cuwedh metpeab anes tec ural Sahe OM, 02. 47. 
Floral leaves up to 4-7.5 mm long 0.75—1.5 mm broad, exceeding 
0) er ee 59. 
Corolla up to 2.3-3.5 mm long, lobes one and a half times as short 
as tube ........ eds soniecpcsuavptbareies curs  seaswsberseeae atoeere tapes eae ieee daeaen dD. 48. 
Corolla up to 4-7 mm long, lobes nearly 2 times or more as short 
AS LUDE iss. -sepesvsseast eave cameacadene nbanicataceinn dad Maree cana ote cae Ra oe 55. 
Flowers mostly on distinct, up to 2 mm long pedicels ........... 49. 


Flowers almost sessile or on extremely short, less distinct pedicels 
wit ae se Remactes a Meas LBs chet alate washec sted aces dagdunat bebaist coataaaumtce aa 50. 


201 


49. 


50. 


5) iB 


Bod 


Ja: 


54. 


3D 


191 


Flowering stems 25—70 cm tall, uniformly sparsely foliated. Corolla 

whitish. Almost entirely herbaceous plant of rubbly steppes in Black 
BEd SUTOUMGINGS ye. cconec cet tccan cee ceesecteeererene- 26. A. rumelica Boiss. 
Flowering stems 20-40 cm tall, more densely foliated in lower 
part. Corolla pink. Subshrub forming lax mats on mountain slopes 
of southern Transcaucasia .................. 47. A. woronowii V. Krecz. 
Flowering stems branched from middle or below, with fairly 
numerous branches. Middle cauline leaves glabrous and smooth 
along margin. Flowers mostly on short, barely distinct pedicels. 
Plant of stony outcrops around Sea Of AZOV .........ceeeeeeeesseeereeeeeeee 
Scectip Sireptenssc Sieur meen te Bh mde tinea 27. A. graniticola Klok. 
Flowering stems branched in upper part, with few (up to 7) branches. 
Middle cauline leaves scabrous-setulose along margin or 
pubescent. Flowers mostly sessile or subsessile. Crimean regions 


Flowering stems scabrous-setulose or pubescent throughout lower 
ones not above 15-20 cm. Plant found in Kerch Peninsula ... 52. 
Flowering stems at top (in inflorescence) glabrous and smooth, 
more developed, up to 25-35 cm tall. Plant found in other regions 
is STB TE, stn ea evtnthane son eet edr iehembcemeteaae at lna nach aha tea D3. 
Flowering stems scabrous-setulose. Cauline leaves scabrous-setulose 
along margin only, short, 3-10 mm long. Caespitose, 5-15 cm tall, 
subshrub, forming small, dense mats ... 35. A. cimmerica V. Krecz. 
Flowering stems pubescent. Cauline leaves 5-15 mm long, similarly 
pubescent. Subshrub 10-20 cm tall, forming larger, lax mats .... 
whee TRS sa tt th te ee 34. A praepilosa V. Krecz. 
Flowering stems densely setose in lower half with distinctly patent, 
0.2-03 or up to 0.5 mm long hairs. Basal and middle leaves with 


Sintilat PUDESCEMCE ..o.-.---seccsene-snceseceecscee est 31. A. praevestita Klok. 
Flowering stems scabrous-setulose in lower part. Leaves setulose 
SEMI PALOH Se ITAD SUN sec ces sect cr cece sesenessscasse rate sncteccuunesasesesse train? 54. 


Cauline leaves 5-18 mm long, about 0.5 mm broad, thin, with 
non-revolute margin, more or less appressed. Floral leaves extremely 
small, 1—-1.5 mm long, 0.2-0.5 mm broad. Corolla 2.5—3 mm long 
sian picts be orale lean uate ca i Roel 32. A. vestita V. Krecz. 
Cauline leaves 3-14 mm long, 0.5-1.5 mm broad, with more or 
less revolute margin, thicker, divaricate from stem almost at right 
angles and arcuately refiexed. Floral leaves 1.5—2.5 mm long, 0.3- 
1 mm broad. Corolla 2-2.5 mm long ......... 33. A. infracta Kiok. 
Corolla glabrous and smooth. Flowering stem up to 30-35 cm tall. 
Transcaucasian Region ...............:.:00+ 48. A. kemulariae Manden. 
Corolla setulose. Flowering stem up to 50-60 cm tall. Plant growing 
in extreme south of European part of the USSR ...............0 56. 


202 


192 


Ti 


58. 


60. 


61. 


Distinctly subshrub ‘with expressed stems ascending from base or 
almost erect, densely velutinus with short almost 0.25 mm long, 
hairs in lower part. Corolla not shorter than 4.5 mm, tube at least 
3 mm long, lobes 2-4 times as short as tube ................:00000 58. 
Almost entirely herbaceous plant with stems ascending or sub- 
erect in lower part, scabrous-setulose. Corolla 3.5-4.5 mm long, 
tube 2.2-2.8 mm long, lobes somewhat less than 2 times as short 
aS tines. AGREE oes vars tanee tes see: ocx coat ruek ad ne a ee ee eee 57: 
Stems smooth above. Middle leaves up to 46 mm long and 1.5 mm 
broad, with smooth margin. Pedicels less distinct, not longer than 
0.5 mm. Corolla 44.5 mm long, with reflexed lobes. Mericarps 
about 2.5 mm long. Plant found in lower Bug Region (vicinity of 
Nilolaev)/..2505 3 Soy ee ee Bees 24. A. hypanica Klok. 
Stems scabrous above. Middle leaves up to 32 mm long, 1.2 mm 
broad, scabrous along margin. Pedicels more distinct, up to 1 mm 
long. Corolla 3.54 mm long, with mildly bent lobes. Fruit 1.5—2 
mm long. Piant found in foothills of Crimea (vicinity of Simferopol) 
5 Sef i, SENSE tryst a Sauk sects 25. A. attenuata Klok. 
Corolla 4.5-4.75 mm long, tube 3—3.5 mm long, lobes 2 times as 
short as tube. Peristome of calyx often with, rarely without, 2 
narrowly linear-filiform, 0.5—0.75 mm long teeth. Plant found in 
the littoral belt of northern Black Sea Coast ............ccsseesseesreeeees 
Ee ee Ret aescusepwesiegy sesssssesseesssesssteeeseeee 23+ A. Didentata Klok. 
Corolla 5.5-7(8) mm long, lobes 2/%2—4 times (usually 2 times) as 
short as tube. Peristome of calyx always without distinct teeth. 
Plant found along coast of Sea of Azov, widely distributed in 


Ce C8 ass: sazasscavatese euyscenct sett eseemets, seetaens 22. A. steveni V. Krecz. 
Flowers on distinct (0.5—2 mm long) pedicels, not aggregated in 
clusters: Entirewflorescence Lax 220.322. casero scsseeseeserreseere n=" 60. 


Flowers sessile or on extremely short, less distinct pedicels, 
aggregated in fairly dense clusters at ends of branches. Entire 
inflorescence more or less CONdENSE ............:.ssccesseeseeeeeeeeeecees 61. 
Flowering stems up to 30 cm tall, glabrous and smooth in upper 
part. Middle cauline leaves narrowly linear to almost setaceous, 
narrowed at apex, up to 30 mm long, with strongly revolute margin. 
Corolla 2.75—3.75 mm long .......... 36. A. biebersteinii V. Krecz. 
Flowering stems 5-8 m tall, sharply scabrous throughout. Middle 
cauline leaves narrowly linear to narrowly linear-spatulate, 
broadened at apex, up to 10 mm long, with almost non-revolute 
margin. Corolla 4.5—-5 mm long ............ 37. A. pedicellata Klok. 
Flowering stems 10-35 cm tall. Corolla 2.5—3.5 mm long ......... 
dhe aphtbe canis caveslvee Mean 2S LE: 2 Re 29. A. lipskyana V. Krecz. 


203 


62. 


63. 


65. 


66. 


193 


Flowering stems not taller than 20 cm. Corolla 4.25-8 mm long 


ate Macrae opaceh ile Mae te oo Soa ta RMR RAR Pty Nae Dal | 62. 
Corolla 3.54.25 mm long, lobes without distinct, whitish, thickened 
tip. Crimean-New Russia low-mountain plant ..................se00 63. 


Corolla not shorter than 4.5-5 mm. Corolla lobes, with distinct, 
often large whitish, thickened tip. Caucasian and Transcaucasian 
Uplands MOscy*alpine Plane oe Lv pene. etoevonsrowsee Mee sasnccsevereeeetnae 64. 

Flowering ‘stems weakly scabrous-setulose in lower part, smooth 
in upper part. Middle cauline leaves setulose only along margin. 

Corolla tube glabrous and smooth or only with isolated fine setae 
at base. Crimean plant ....................4 30. A. aemulans V. Krecz. 
Flowering stems pubescent throughout (hairs about 0.2 mm long), 
middle cauline leaves densely pubescent with patent hairs up to 
0.3 mm long on both surfaces. Corolla tube pubescent. Plant found 
in Novorossiisk area ............scssesseeeeeseseees 28. A. markothensis Klok. 
Flowering stems fairly densely puberulent throughout, with separate, 
thin, extremely fine hairs (not setae!). Leaves similarly puberulent 
along margin, the midrib above and beneath. Corolla tube puberulent 
throughout, lobes barbate with more distinct hairs along margin at 
apex. Flowers in terminal 3-flowered clusters on shortened branches 
of inflorescence. Plant found in alpine areas of Ingushetia ......... 
NS Sertaas La Same tenetet eee eto areateots ed 38. A. dazyantha Klok. 
Flowering stems scabrous-setulose, scabrous or smooth. Leaves 
scabrous-setulose only along margin, rarely weakly scabrous also 
on surface. Corolla tube with scattered setae only at base or entirely 
smooth and glabrous; lobes glabrous and smooth. Flower clusters 
FINA MO WET Rie. cee sorter. .eSeeetres nets dace satereen cocthos cecteeee tector actoren 65. 
Floral leaves at base of floral clusters up to 4-5 mm long, 
considerably shorter than flowers. Upper cauline leaves shorter 
than internodes. Anthers oblong, 1-1.25 mm long. Tubercles on 
ovary and mericarps obtuse, almost hemispherical. Plant found in 
Ciscaucasia, Dagestan and western Transcaucasia ...............+. 66. 
Floral leaves at base of floral clusters 3-7.5 mm long, exceeding 
or almost equaling flowers. All cauline leaves longer than 
internodes. Anthers linear, about 1.5 mm long. Tubercles on ovary 
and mericarps mucronate, cuneate. Iranian plants, extending to 
SOMME FMP NANSCAUCASI Ale sei 5525 TEES eactese oath ootenesene ne 71. 
Flowering stems 10-20 cm tall, weakly scabrous due to retrorse, 

and almost unnoticeable, subacute, cuneate papillae throughout. 

Middle cauline leaves 10-20 mm long, 1—2 mm broad; floral leaves 
linear or oblong-lanceolate, up to 5 mm long and 1.5 mm broad. 
Corolla sao MMT LONE es... cesccttencesoteee 39. A. accrescens Klok. 


204 


194 


67. 


68. 


69. 


70. 


iL, 


ie 


73. 


Flowering stems up to 10 or 15 cm tall, scabrous-setulose, acutely 
scabrous or smooth, but without retrorse papillae. Middle cauline 
leaves up to 10 mm rarely up to 17 mm long, and not broader than 
1 mm; floral leaves either considerably narrower or just as broad, 
but ovate-lanceolate. Corolla 5—6 mm, rarely up to 7 mm long .... 
Ee EARN LAE LM Ne a eS 67. 
Flowering stems glabrous and smooth throughout. Leaves smooth 
over entire margin. Corolla tube entirely smooth .................0000 
RNS RUT BUEN VEE a ES 42. A. intersita Klok. 
Flowering stems scabrous-setulose throughout or at least in lower 
part and above, below nodes. Cauline leaves scabrous-setulose along 
entire margin, or only in lower part. Corolla tube with scattered 
setae at base, rarely smooth all Over ...............::cssccsessesseeereeeeees 68. 
Cauline leaves linear or oblong-lanceolate, narrow, 0.25—0.75 mm 
broad. Flowers often in clusters of 3, rarely 5. Plant found in 


Ciscaucasia..and Dagestan, anc. ctegin fansite Deep cet nitpeteestcye et. sti 69. 
Floral leaves oblong or ovate-lanceolate, 0.5—1.5 mm broad. Flowers 
in clusters of 3-9. Plant found in western Transcaucasia ....... 70. 


Flowering stems 5—10 cm tall. Cauline leaves from narrowly linear- 
spatulate (basal) to linear-lanceolate, flat, 0.5—1.25 mm broad. Entire 
plant bright green ..... 40. A cristata (Somm. and Lev.) V. Krecz. 
Flowering stems mostly 10-15 cm tall, slender. Cauline leaves 
narrowly linear, with more or less revolute margin, 0.5—0.75 mm 
broad. Grayish-green plant ............:cssescssseseseeeeeees 41. A. alpina M.B. 
Cauline leaves up to 12 mm long, almost flat; floral leaves setulose 
along margin and apex. Corolla tube with scattered setae in lower 
part. Dark green plant, blackish when dry....44. A. pontica Boiss. 
Cauline leaves up to 17 mm long, with more or less revolute margin. 
Radical leaves glabrous and smooth along margin. Corolla tube 
entirely glabrous and smooth. Grayish-green plant, not blackish 
NOTE Visi a a nt 43. A. abchasica V. Krecz. 
Cauline leaves 3-12 mm long, 0.25-0.75 mm broad; floral leaves 
longer than flowers. Corolla 5-5.5 mm long. Style 1-1.5 mm long 
EL OE ena Viele» 45. A. affinis Boiss. and Huet. 
Cauline leaves 7-18 mm long, 0.5—1 mm broad. Floral leaves 
somewhat shorter than flowers. Corolla 5.5-6.5 mm long. Style 
WS VID 52a, ah coe nt cece ees 46. A. dolichophylla Klok. 
Corolla lobes equaling or almost equaling tube ...................0. 73: 
Corolla lobes considerably (142-2% times) shorter than tube ...77. 
Flowering stems 35-50 cm tall, ascending from base in lower part, 
densely covered by distinctly noticeable setaceous hairs together 
with leaves, upper part glabrous and smooth or weakly scabrous. 


205 


Plant found along northern coast of Black Sea ..........ccccssessseseeeesseees 

bee Mme ED AO, EL RIE. cass wostenersaveesenes 49. A. setulosa Boiss. 

+ Flowering stems 10-35 cm tall, usually obliquely ascending; in 
lower part scabrous-setulose or glabrous, above glabrous and 
smooth, rarely setulose. Plant found in different regions ........ 74. 

74. Fruit (and ovary) without or with scattered scales. Plant endemic 
for sand of middle reaches of Dmieper ...............:ccsceseeseseeseeeeees 

UB ee | UD Seba. 52. A. leiograveolens M. Pop. and Chrshan. 

‘Pp wrerruit densely covered with scales Wil. SA is. 
75. Flowering stems scabrous-setulose up to the tip, together with leaves. 
Corolla 3—3.5 mm long. Fruit 1.75-3 mm long. Plant endemic for 

sands of southern Bug and Savran on the right banks in Ukraine 

TRIBE BU ROR RNS AEROS BE 50. A savranica Klok. 
+ Flowering stems glabrous and smooth in upper part and upper 
leaves. Corolla 3.54.5 mm long. Fruit 34.5 mm long. Plant found 
imLother more Caster Tesions eee a EAN BL ee 76. 

76. Flowering stems scabrous-setulose in lower part, together with 
leaves. Cauline leaves up to 40 mm long. Plant found in riverside 

sands between Dnieper and Volga .......... 51. A. graveolens M.B. 

+ Flowering stems glabrous and smooth to very base. Cauline leaves 
up to 2.5 mm long, indistinctly scabrous only along margin. Plant 

found on coast of Caspian Sea in plains of Dagestan ................. 
ene rs Tune he ea BO ee 53. A. diminuta Klok. 
77. | Flowering stems scabrous in lower part. Middle cauline leaves 10— 
20(24) mm long, several times as short as internodes, distinctly 
connate at base (equaling width of leaf). Corolla lobes 2—2% times 

as short as tube. Plant found in Aralo-Caspian seas region 

Oped DON i VRE BI Bh Ola. Cie 55. A. danilewskiana Basin. 

+ Flowering stems smooth up to base. Middle cauline leaves up to 40 
cm long, one and a half two times as short as internodes, indistinctly 
connate at base. Corolla lobes 142-2 times as short as tube. Plant 
found in south-eastern regions of European part of USSR 

JE PGC RSE BGP ea Een Petes, 54. A. laevissima Klok. 


Section 1. CrucIANA Gris. Spicil. fl. rumel. IT (1844) 166; Boiss. 
Fl. or. II, 25; Schumann in Pflanzenfam. IV, 4, 5, 148.— Flowers in 
spicate or racemose, pseudowhorls, five-merous. Corolla with long 
tube, 3-3% times as long as lobes, opening in evening with lobes 
having incurved tips. Leaves in whorls of 6-14. 


Series 1. Molluginoideae Pobed.—Inflorescence spicate, consisting 
of clusters of flowers, aggregated in head. Leaves in whorls of 8-14, 
large. 


206 


196 


This series also includes A. kotschyana Boiss. et Hohen., described 
from Kurdistan. 


1. A. molluginoides (M.B.) Rchb. Fl. Germ. exs. I (1830-1832) 
206; Boiss. Fl. or. III, 27; Schmalh. Fl. II, 8; Javorka, Magyar fl. III, 
1036(?); Grossh. Fl. Kavk. IV, 22.—Crucianella molluginoides M.B. 
Fl. taur.-cauc. I (1808) 106 and III (1891) 110; DC. Prodr. IV, 587; 
Ldb. Fl. Ross. II, 404. —Ic.: M.B. Cent. pl. rar. ross. I, tab. 65. 

Perennial. Rootstock thick, woody, almost horizontal, red-dark 
brown. Stems several, 30—80(100) cm tall, straight, simple, rarely 
weakly branched, with short lateral twigs, extremely rarely densely 
branched, branched long, foliated almost from the very base, 8-sided, 
glabrous and retrorsely pubescent along ridges, with dense, short 
pubescence at nodes and along ridges. Leaves in distant whorls of 8— 
14, oblong-linear, (22)35—50(57) mm long, (1)2—2.5(4.5) mm broad, 
gradually acuminate at apex, ending in short cusp, with short pubescent 
ring at base, with one sharply protruding vein, involuted along margin; 
antrorsely pubescent along margin as well as along midrib beneath. 
Floral leaves in whorls of 8—10, much smaller. Bracts ovate or elliptic, 
(2)3—4(5) mm long, (1)1.5(2) mm bread, with cusp at apex, with long, 
white cilia along margin longer than width of bracts, rarely, 
predominantly in southern Transcaucasia, with short cilia, just like 
leaves (var. breviciliata Pobed.). Flowers in sessile clusters, aggregated 
in head on peduncles pubescent along ribs forming narrow, racemose, 
interrupted or sometimes almost paniculate inflorescence in upper stems 
and branches. Peduncles in upper heads extremely short, successively 
longer in lower whorls of leaves. Corolla with very long tube, 6—12 
times as long as lobes; lobes 5, ovate, sharply narrowed at apex into 
thick, long inflexed tip, 2 mm long including tip. Stamens 5, with 
filaments considerably. shorter than anthers; anthers 2 mm long, not 
exserted from corolla tube. Style longer than half of corolla tube, 
bipartite above middle, with unequal lobes, with thick hypogynous 
disk at base in form of small collar; stigma subglobose. Fruit binate; 
mericarps oblong, 2.5 mm long, 1.5 mm broad, brown, strongly 
granular. May to August. 

On mountain-meadow slopes, in subalpine zone, extending to 2000 
m.— Caucasus: Ciscaucasia, eastern and southern Transcaucasia. 
General distribution: Armenia-Kurdistan. Described from Kislovodsk. 
Type in Leningrad. 

Note. The locality of this species in Hungary, reported by 
Reichenbach (Rchb. l.c.), is doubtful. Judging from the drawing, 
provided by Reichenbach (Rchb. Ic. Fl. Germ. XVII, tab. 1176), it, 


207 


197 


evidently is another species. Javorka (l.c.) also doubts the occurrence 
of A. molluginoides in Hungary. 


Series 2. Prostratae Pobed.—Inflorescence spicate-paniculate, 
clusters of flowers opposite, interrupted. Leaves in whorls of 6, green 
with mildly involuted margin. 


2. A. prostrata (Adams) C. Koch in Linnaea, XXIV (1851) 461; 
Grossh. Fl. Kavk. IV, 23.—A. aspera (M.B.) Boiss. Fl. or. III (1857) 
28.—Crucianella prostrata Adams in Weber et Mohr, Beitr. I (1805) 
46.—C. aspera M.B. FI. taur.-cauc. I (1808) 107; DC. Prodr. IV, 588; 
Ldb. FI. Ross. II, 404.—A. aspera var. latifolia Boiss. Fl. or. III (1857) 
28 p.p.—A. prostrata var. pubiflora Bordz. in Visn. Kiivs’k. bot. sadu, 
XII-XII (1931) 139. 

Perennial. Root thick, vertical, woody, light-red-brown. Stems 
numerous, procumbent at base, ascending, 10-40 cm tall, simple or 
branched mainly in lower part, with long branches reaching tip of 
plant, terete at base, whereas 4-sided above and in branches, densely 
pubescent. Leaves in whorls of 6, narrowly linear, (9)13—15(18) mm 
long, 0.5—1 mm broad, with short cusp at apex involuted and scabrous 
due to short, upward directed setae along margin, with single vein. 
Floral leaves smaller. Flowers in opposite, interrupted clusters, upper 
clusters sessile, lower on more or less long, pubescent peduncles, 
forming narrow spicate-paniculate, inflorescence. Bracts lanceolate or 
elliptic, 2-4 mm long, 0.75-1.5 mm broad, with long thin cusp at 
apex, long ciliate along margin especially in lower part, 2 times as 
short as corolla. Corolla with long glabrous or pubescent (var. 
pubiflorum Bordz.) tube, 3-3% times as long as lobes; lobes 5, oblong, 
2 mm long (taking into account inflexed tip), apex with thick inflexed 
tip almost equaling lobe. Stamens 5, on extremely short filaments, 
attached at middle of long, narrow anthers. Style bipartite from middle 
or above with identical lobes; stigma hemispherical. Fruit binate; 
mericarps oblong, 2 mm long, microgranulate. June to August. 

On limestone, stony and rubbly slopes, rocks.— Caucasus: Western 
Transcaucasia (only southern part), eastern and southern Transcaucasia, 
Talysh. General distribution: Armenia-Kurdistan, Iran-Afghanistan 
(northern part). Described from Ossetia. Isotype in Leningrad. 


3. A. albovii Mand. in Grossh. Opred. rast. Kavk. (1949) 247.— 
A. aspera (M.B.) Boiss. var. latifolia Boiss. Fl. or. III (1857) 28 
p-p.—lIc.: Fl. Gruzii, VIII, fig. 359. 


208 


198 


Perennial. Root thick, long, vertical, woody, dark red-brown or 
gray-reddish brown. Stems numerous, procumbent at base, ascending, 
40-50 (rarely 10) cm tall, more branched than in the foregoing species, 
with numerous, predominantly sterile branches at base, 4-sided from 
base and densely pubescent along ridges or terete at base and pubescent 
over entire surface, 4-sided in remaining part and in branches and 
pubescent only along ridges. Leaves in whorls of 6, linear or narrowly 
elliptic, (5)7—8(11) mm long, (0.5)1.5(2) mm broad, with short cusp 
at apex, slightly involuted and scabrous due to short, upward directed 
setae along margin, sometimes with similar pubescence on upper 
surface, with single vein. Flowers in opposite, lax, few-flowered, 
interrupted clusters, upper clusters sessile, lower on more or less long, 
pubescent peduncles, forming spicate-campanulate inflorescence. Bracts 
predominantly lanceolate or elliptic, (2.5)3(4) mm long, (0.5)1(2) mm 
broad, with long thin cusp at apex, long ciliate along margin, especially 
in lower part, 2 times as short as corolla. Corolla with long, glabrous 
tube, 3—-3% times as long as lobes; lobes 5, oblong, 2 mm long 
(including inflexed tip), apex with thick inflexed tip almost equaling 
lobe. Stamens 5, on extremely short filaments, attached to middle of 
long narrow anthers. Style bipartite from middle or above, with unequal 
lobes; stigma subglobose. Fruit binate; mericarps oblong, 2-3 mm 
long, microgranulate. June to August. 

On rubbly and stony slopes, rocks, screes, in alpine and subalpine 
zones.— Caucasus: Ciscaucasia, Dagestan, western Transcaucasia. 
Endemic. Described from Dagestan (Andi Range). Type and isotype 
in Leningrad. 

Note. A. albovii is a good geographical race with its range almost 
completely separated from that of A. prostrata. A. albovii is widespread 
over the Greater Caucasian Range, whereas A. prostrata is widespread 
over Lesser (?) Caucasian Range and its ridges, and extends to Iran 
and Asia Minor. Both species are sympatric only on southern slopes of 
the Great Caucasian Range in South Ossetia. 


Series 3. Glomeratae Pobed.—Inflorescence spicate, interrupted, 
rarely almost continuous. Leaves in whorls of 4-6-7, fleshy, grayish, 
involuted along margin from midrib. 

This series also includes these closely related species from outside 
the USSR: A. capitata (Zabell.) Grossh. from Lebanon and northern 
Iran, and A. bracteata (Boiss.) Grossh. from northern Iran. 


4. A. glomerata (M.B.) Griseb. Spicil. fl. rumel. IT (1844) 166; 
Boiss. FI. or. III, 28; Grossh. Fl. Kavk. IV, 23. —Crucianella glomerata 


21 


— 


199 


M.B. FI. taur.-cauc. I (1808) 107 and III (1819) 110; DC. Prodr. IV, 
588; Ldb. Fl. Ross. II, 404.—Exs.: Herb. fl. Cauc. No. 390. 

Perennial. Root thick, woody, reddish. Stems numerous, 7-35 cm 
tall, straight or ascending, branched with short densely foliated sterile 
branches, terete, densely pubescent with long patent hairs. Leaves in 
whorls of 6, rarely 7, linear, (4)7-9(10) mm long, 0.5—1 mm broad, 
short acuminate, sessile, strongly involuted along margin, with one 
vein, fleshy, grayish, pubescent above with short, rigid upright setae, 
lower old leaves declinate, appressed to stem, floral leaves smaller. 
Bracts lanceolate or ovate-lanceolate, (5)6-7 mm long, (1)2(3) mm 
broad, half or one third as short, sometimes 2—3, connate up to one 
third of length along margin, shortly, and in lower half usually long 
ciliate, pale green. Flower clusters opposite, many-flowered, sessile, 
forming spicate, interrupted inflorescence at ends of stems and 
branches. Corolla greenish, with long, glabrous or sparsely pubescent 
tube, 2 times as long as glabrous lobes, lobes oblong or ovate-oblong, 
2 mm long (including inflexed tip), apex with thick inflexed tip half 
as long as lobe. Stamens 5, anthers narrow, long; filaments short, 
attached to middle of anthers, scarcely exserted from throat of flower. 
Style bipartite above middle, with unequal lobes, reaching as far as 
middle of corolla tube or higher. Stigma globose. Fruit binate; mericarps 
oblong, 2.5-3 mm long, 1-1.5 mm broad, extremely microgranulate, 
glabrous or covered with white hairs. June to August. 

In dry, stony places, rocks, up to middle mountain zone.— 
Caucasus: Dagestan, eastern and southern Transcaucasia, Talysh. 
General distribution: Armenia-Kurdistan, Iran-Afghanistan (north). 
Described from Georgia (vicinity of Tbilisi). Type in Leningrad. 


5. A. hirsutiuscula Pobed. sp. n. in Addenda XXII (1958) 690. 

Perennial. Root thick, woody, reddish. Stems numerous, 10-20 
cm tall, straight or ascending, branched, with short densely foliated 
sterile branches, terete, scabrous due to extremely short, dense setae. 
Leaves in whorls of 6-8, linear, (4)7(8) mm long, (0.5)1(1.25) mm 
broad, short-acuminate sessile, strongly involuted along margin, with 
single vein, fleshy, grayish, pubescent above with short rigid upright 
setae, lower old leaves declinate, appressed to stem. Floral leaves 
smaller. Bracts broadly elliptical or broadly ovate, 5-7 mm long, 4— 
6 mm broad, one per flower, equaling flowers or somewhat longer, 
scabrous on both surfaces due to dense, extremely short setae, besides 
two extremely narrow, small, linear bracts with similar pubescence. 
Flower clusters opposite, many-flowered, sessile, forming spicate, 
interrupted inflorescence at ends of stems and branches. Corolla yellow 
(recorded in label of color of live corolla), with long, setulose tube 


209 


200 


Wy), 


Deen Ty . 
. Mari ey 


"MA AN Ap, A Pearly wry, 
RENN i oe 
Se) = | : 


J 

\ 

Sl Ne i2 
A YW 


Plate XI. 
1. Asperula caxcasica Pobed., upper part of plant, unfolded corolla, style, 
mericarps.— 2. A. taurica Pacz., general appearance of plant, unfolded corolla, 
pistil, coccus.—3. A. balchanica Bobr., general appearance of plant, unfolded 
corolla, pistil—_ 4. A. ferganica Pobed., general appearance of plant, unfolded 
corolla, pistil, coccus. 


212 


201 


4—5 times as long as similarly pubescent lobes; lobes broadly ovate, 
0.5 mm long, apex with thick, long, incurved tip. Stamens 5. Anthers 
narrow, long; filament several times as short as anther, attached towards 
its middle scarcely exserted from throat of flower. Style, bipartite 
about the middle with unequal lobes, reaching as far as middle of 
corolla tube or higher. Young fruit binate, densely microgranulate, 
diffusely setulose, covered with white bands. May. 

On slopes of clayey hills. — Caucasus: Eastern Transcaucasia. 
Described from Kobystan (Perekesh-Kul). Type in Leningrad. 

Note. It is closest to A. bracteata (Boiss.) Grossh. of northern 
Iran, from which it differs by broader short-setose bracts without long 
cilia on the petioles and by broader short-setose pubescence on the 
stem, corolla and fruits. 


6. A. ferganica Pobed. sp. n. in Addenda XXII, 690.—A. 
glomerata auct. fl. As. med. non Griseb.: O. and B. Fedch. Perech. 
rast. Turkest. III (1909) 135, p.p. 

Perennial. Root thick, woody, vertical, light reddish. Stems 
numerous, 10-40 cm long, straight or ascending, with short densely 
foliated sterile branches, terete, densely pubescent with short, appressed 
hairs. Leaves in whorls of 6, rarely 7, linear, (S)10—12(18) mm long, 
(0.5)1(1.5) mm broad, short-acuminate, sessile, strongly involuted along 
margin, with single vein, fleshy, grayish, pubescent above with short 
rigid upright setae, lower old leaves declinate, floral leaves smaller. 
Bracts lanceolate or ovate-lanceolate, (4)10(11) mm long, (1.5)2.5(4) 
mm broad, one per flower, surrounding ovary and, besides, 2 per 
flower, linear, one-third length of flowers, connate up to middle or 
one third, along margin short, in lower half usually long ciliate, in 
lower part almost flat, shorter than flowers. Flower clusters opposite, 
multi-flowered, sessile forming, spicate, interrupted inflorescence at 
ends of stems and branches. Corolla purple, with long tube, gradually 
spreading above, glabrous, 5S—6 times as long as glabrous lobes, lobes 
5, lanceolate or oblong-ovate, 1.5 mm long (including incurved tip), 
apex with thick tip incurved up to half the length of lobe. Stamens 5; 
anthers narrow, long; filaments 4 times as short as anthers, inserted in 
corolla tube much below lobes. Style with 2 unequal lobes, above half 
length, reaching as far as 3/4 length of corolla tube; stigma globose. 
Fruit binate; mericarps oblong, 3-4 mm long, 1.5-2 mm broad, 
microgranulate, with extremely fine papillae on surface, sometimes 
with white bands. Flowering May to July. Fruiting August to September 
(Plate XI, fig. 4). 


21 


Ww 


202 


On stony slopes, shingly screes.—Soviet Central Asia: Pamiro- 
Alai (northwestern part), Tien Shan. Endemic. Described from Fergana 
(Gulcha). Type in Leningrad. 


7. A. pamirica Pobed. sp. n. in Addenda XXII, 691.—A. glomerata 
auct. fl. As. med. non griseb.: O. and B. Fedch. Perech. rast. Turkest. 
(1909) 135, p.p. 

Perennial. Root extremely thick, woody, vertical, light reddish, 
with short, thick, large caudexes at top. Stems numerous, 10-25 cm 
tall, straight or ascending, with short densely foliated sterile branches, 
terete, densely pubescent with more or less long, patent hairs. Leaves 
in whorls of 8 on main stem and in whorls of 6 on branches, linear, 
(5)8(12) mm long, (0.5)1(1.5) mm broad, short-acuminate, sessile, 
strongly involuted along margin, with single vein, fleshy, grayish, 
pubescent above with short rigid upright setae, lower old leaves 
declinate; floral leaves below inflorescence almost similar to cauline 
leaves. Bracts almost equaling flowers, lanceolate-ovate, one per flower, 
surrounding ovary, besides two linear bracts, occasionally connate to 
one third or half setulose in upper part, with long silky hairs in 
remaining part and along margin, making entire involucre white- 
pubescent. Flower clusters opposite, many-flowered, sessile, forming 
spicate, interrupted inflorescence at end of stems and branches. Corolla 
yellowish-greenish, with long tube, 5 times as long as lobes, not 
spreading above, densely pubescent on outside, apex of lobes having 
a cluster of long upright, straight hairs at curvature; lobes broadly 
ovate or ovate, with thick, herbaceous incurved tip, equaling almost 
half of lobe. Stamen 5, with short filaments, shorter than anthers, 
inserted in corolla tube below lobes; anthers barely visible in throat. 
Style bipartite from base or from middle; stigma subglobose. Fruit 
binate, with unequal mericarps, one rudimentary, 3 mm long, 1.5 mm 
broad, glabrous, microgranulate on surface, with extremely fine papillae 
and white bands. Flowering June. Fruiting July to October. 

Mountain steppe slopes, screes in upper mountain belt.— Soviet 
Central Asia: Pamiro-Alai. Endemic. Described from Shugnan. Type 
in Leningrad. 

Note. Attention must be called to the specimens from Daraut- 
Kurgan, which have characters of both species.—A. ferganica and A. 
pamirica; in the pubescence of the flowers it is A. pamirica, but the 
flowers have a distinct purple color as in A. ferganica, and the stems 
have the latter’s characteristic short pubescence. Evidently, in this 
region, these species produce hybrids. 


214 


203 


8. A. balchanica Bobr. in Tr. Bot. sada, XLIV (1931) 74; descr. 
in Addenda XII, 691. 

Perennial. Root extremely thick, woody, light brown, multiheaded. 
Stems numerous, extremely short, 1.5—7 cm tall, branched, forming a 
dense mat, terete, puberulent appressed or barely patent curved hairs. 
Leaves usually in whorls of 4, rarely 6, extremely small, linear, 2-3 
mm long, 0.5—0.75 mm broad, very thick, with highly involuted margin, 
densely covered with thick setulae, with short, strong cusp at apex, 
single veined, sessile; floral leaves below inflorescence in threes, 
differing only slightly from cauline leaves. Bracts broadly ovate one 
per flower besides 2 narrow linear bracts several times as short as 
flowers, patently pubescent along margin, and upper part with clusters 
of long, white, thin hairs at base. Flowering stems hardly rising above 
dense pulvinate vegetative shoots. Flowers in few-flowered (2-4) 
clusters at ends of stems and branches, sessile. Corolla purple, with 
long tube and short lobes; lobes with almost as long fleshy tip, inflexed, 
puberulent outside, but without clusters of long, straight hairs on fold 
of tip. Stamens on extremely short filaments, several times as short as 
anthers, inserted below lobes, with anthers almost up to half exserted 
from throat of flower. Style reaching middle of corolla tube, bifid 
from middle, with globose stigma. Ovary diffusely puberulent, 
microgranulate and, besides, covered with numerous short white hairs. 
Fruit unknown. June (Plate XI, fig. 3). 

On stony slopes and mountain peaks.—Soviet Central Asia: 
mountainous Turkmenia (only in Great Balkhans). Endemic. Described 
from Great Balkhan Mountains). Type in Leningrad. 


9. A. turcomanica Pobed. sp. n. in Addenda XII, 692.—A. 
glomerata auct. non Griseb.: O. and B. Fedch. Perech. rast. Turkest. 
III (1909) 135.—Exs.: Ed. Hort. Bot. Petr. Magni, No. 95. 

Perennial. Root thick, woody, light reddish brown, with small 
caudex. Stems numerous, straight or ascending, 10—35 cm tall, with 
long erect, terete lateral branches, densely pubescent with long, patent 
hairs. Leaves in distant whorls of 6, occasionally in closely spaced 
whorls of 4 on branches, linear, (4)7—8(11) mm long, (0.25)0.5(1) 
mm broad, extremely thick with highly involuted margin reaching 
middle vein, densely covered with thick setulae, having short cusp at 
apex, single veined, sessile; floral leaves below inflorescence in whorls 
of 4—5, differing slightly from cauline leaves. Bracts lanceolate, narrow, 
long, somewhat shorter or almost equaling flowers, (6)8—10(15) mm 
long, (1)2.5(3) mm broad (one per flower) and linear, narrow (2 per 
flower), all narrowly and long tapering at apex, with long, white cilia 
along margin up to apex, setulose on both surfaces. Inflorescence 


215 


204 


spicate at ends of stems and branches, comprising closely spread 
opposite clusters. Corolla (purple?) puberulent outside, with 5 mm 
long tube; lobes short, ovate, 1.5 mm long, with thick tip, incurved 
almost up to middle of lobe. Stamens with extremely short filaments, 
inserted in corolla tube below lobes, anthers long, up to half exserted. 
Style extremely long, bifid from middle, upright from flower or 
reaching only up to half length of tube. Fruit with or without diffuse 
puberulence, covered with abundant short white hairs. April to June. 

On stony and rubbly slopes of mountains, screes.— Soviet Central 
Asia: mountainous Turkmenia. General distribution: Iran-Afghanistan 
(Astrabad province). Described from Kopet-Dag (Ioldere Gorge). Type 
in Leningrad. 


Section 2. MONOCEPHALA Pobed. sect. n. in Addenda XXII, 692.— 
Sect. Cynanchicae DC. Prodr. IV (1830) 582, p.p.—Sect. Cynanchica 
§ latifoliae Boiss. Fl. or. III (1875) 26; Schumann in Pflanzenfam. IV 
abt. 4 and 5, 119.—Inflorescence single-headed at ends of stems and 
branches, covered by involucre leaves. Flowers 4-merous, with 
extremely long corolla tube, 3—4 times as long as lobes. Leaves in 
whorls of 4. 

Besides our species, two western European species belong to this 
section: A. taurina L., widespread in southern Europe from Spain to 
Hungary, and A. leucanthera Beck., growing in the Balkan Peninsula, 
where A. taurina is not found, and in southern Austria, where both 
species occur together. Moreover, this section contains two species 
from Asia Minor: A. involucrata Walhen. and A. suavis Fisch et Mey 
(Turkish Armenia). 


10. A. caucasica Pobed. in Addenda XXII, 692.—A. taurina auct. 
non L.: M.B. FI. taur.-cauc. I, 100; DC. Prodr. IV, 582, p.p.; Ldb. FI. 
Ross. II, 397, p.p.; Boiss. Fl. or. III, 41, p.p.; Schmalh. Fl. II, 7, p.p.; 
Grossh. Fl. Kavk. IV, 24.—Exs.: Herbarium of the Russian Flora, No. 
LHF: 

Perennial. Rootstock creeping, woody, reddish. Stem straight or 
ascending, 20-80 cm tall, simple or branched with long branches, 
sometimes reaching top of stem, 4-angled, glabrous or diffusely 
pubescent with patent hairs. Leaves in whorls 4, ovate-lanceolate or 
broadly elliptical, (20)50-—60(80) mm long, (10)20—25(30) mm broad, 
narrowed or gradually long tapering toward apex, without cusp, at 
base abruptly narrowed into extremely short petiole or almost sessile, 
entire, with 3 or 5 veins, stiff scattered pubescence on both surfaces, 
densely pubescent along veins and margins paler beneath. Inflorescence 
single-headed, terminal on stem and branch, many-flowered, surrounded 


216 


205 


by involucre leaves, outer ones lanceolate, resembling leaves, but 
smaller, inner ones linear, narrow, (5)7(10) mm long, 1—1.5 mm broad, 
long ciliate along margin (cilia several times as long as leaf breadth), 
about half as short as flowers. Flowers mixed, bisexual, or staminate. 
Corolla white, rarely pink, with long, almost filiform 8-9 mm long 
tube extremely narrow at base, densely, appressed puberulent, 
infundibuliform, with 4 linear, 2-3 mm long, obtuse lobes. Stamens 
on long filaments, almost equaling anthers or somewhat shorter, with 
narrow, violet, 1.5—2 mm long anthers, exserted from flower. Style 
bifid at tip (last quarter length); stigma capitate or almost globose. 
Fruit binate; mericarps almost globose, 4 mm long, and broad, giabrous, 
microgranulate, and rugose. April to August (Plate XI, fig. 1). 

In beech and other broad-leaved forests, scrub.— European USSR: 
Crimea (very rarely and not entirely convincingly). Caucasia: 
Ciscaucasia, western, eastern and southern Transcaucasia. General 
distribution: Armenia-Kurdistan, Iran. 

Described from Abkhazia (Tsebelda). Type in Leningrad. 

Note. This species closely resembles the Western European species 
A. taurina L., which it was taken to be earlier, but it differs from it 
in having longer and denser pubescence along the veins and scattered 
hairs over the entire surface on both sides of the leaves, densely long- 
ciliate inner bracts of the involucre, often five veins on the leaves, 
somewhat shorter corolla lobes, ca. 2-3 mm long, shorter stamen 
filaments. 


11. A. propinqua Pobed. in Addenda XXII, 693.—A. taurina 
auct. non L.: Ldb. Fl. Ross. II, 397, p.p.; Boiss. Fl. or. III, 41, p.p.; 
Schmalh. Fl. III, 7, p.p.—A. leucanthera auct. non Beck: Stank. and 
Tal. Opred. (1949) 577. 

Perennial. Rootstock creeping, woody, reddish. Stem straight or 
rising, 30-50 cm tall, simple or branched with long branches, 
sometimes reaching top of stem, 4-angled, glabrous or with scattered 
patent hairs, especially in upper part. Leaves in whorls of 4, ovate- 
lanceolate or broadly elliptical, (30)40-45(52) mm long, (12)13—14(17) 
mm broad, attenuated at apex, without cusp, at base suddenly narrowed 
into short petiole or almost sessile, entire, with 3 veins, diffusely 
pubescent on both surfaces, densely pubescent along veins and margin, 
paler beneath. Inflorescence single-headed, terminal on stem and 
branch, many-flowered, surrounded by involucre leaves, outer ones 
lanceolate, resembling leaves but smaller, inner linear, narrow, (5)6— 
7(8) mm long, (0.5)1(1.5) mm broad, long ciliate along margin (with 
cilia several times as long as breadth of leaves), about half as short as 
flowers. Flowers bisexual and staminate. Corolla white, 6-8 mm long 


217 


206 


almost filiform with tube, extremely narrow at base, densely appressed 
puberulent together with lobes, infundibuliform; lobes 4, linear, 2—3 
mm long, obtuse. Stamens on long filaments, almost equaling anthers 
or shorter, with narrow, white (light-brown in herbarium) 1.5—2 mm 
long anthers, exserted from flower..Style included in tube, bifid at tip, 
with identical lobes; stigma capitate or almost globose. Fruit binate, 
easily dividing; mericarps almost globose, 3 mm long and 2 mm broad, 
microgranulate and rugose. April to July. 

In forests on mountain slopes.— European USSR: Crimea. 
Endemic. Described from Mt. Agarmysh. Type in Leningrad. 

Note. Closely resembling the Western European species A. 
leucanthera Beck., it differs in having leaves that are more attenuated 
at the apex, pubescent with scattered hairs over the entire surface 
(denser and longer along veins), somewhat shorter corolla lobes, 2.5— 
3 mm long, short stamen filaments, equaling or shorter than anthers. 
The separation of this species from the even closer Caucasian species 
A. caucasica Pobed. is the same as the separation of A. leucanthera 
Beck. from A. taurina L., i.e., it is distinguished by its white anthers 
and some other smaller characters. 


Section 3. TRICHOCARPAE Pobed. sect. n. in Addenda XXII, 694.— 
Inflorescence terminal, with 3 simple or furcate or ternately branched 
cymes. Flowers 4-merous, with short corolla tube, shorter or equaling 
lobes. Leaves in whorls of 4—10. 


12. A. odorata L. Sp. pl. (1753) 103; M.B. FI. taur.-cauc. 1, 100; 
Ldb. FI. alt. I, 139; DC. Prodr. IV, 585; Ldb. Fl. Ross. II, 400; Boiss. 
FI. or. III, 43; Schmalh. Fl. II, 8; Kom. and Alis. Opred. rast. Dalnevost. 
kr. II, 947; Kryl. Fl. Zap. Sib. X, 2571; Grossh. Fl. Kavk. IV, 24.— 
Ic.: Syreistsch. Fl. Mosk. gub. III, 187; Fedch. and Fler. Fl. 906; Hegi, 
Ill. Fl. VI, 1, tab. 247.—Exs.: Fl. Gall. et Germ. exs. No. 1493; FI. 
Boh. et Morav. exs. No. 1087; Kickxia Belgica, No. 321; Fl. exs. 
Reipubl. Boh.-Slov. No. 273; Pl. D’Espagne, No. 6580; FI. stir. exs. 
No. 144; FI. lit. exs. No. 78; Pl. Finl. Exs. No. 1349; Eston. pl. no. 
176; Fl. Pol. exs. No. 177; GRF, No. 880. 

Perennial. Rootstock small, creeping, branched. Stem single, 
straight, simple, 10-60 cm tall, 4-angled, smooth, glabrous. Leaves in 
whorls of 6-10, broadly oblanceolate, (18)40(54) mm long, (4)10(13) 
mm broad, with short cusp at apex, narrowed at base, antrorsely 
setulose along margin and sometimes along vein beneath, or diffusely 
appressed setulose on both surfaces, with a ring of short setae at base 
of leaves; floral leaves 4—6 at base of inflorescence and in pairs on 
their branches, the latter extremely small, (1)1.5-2(4) mm long, 


207 


0.25-—0.5 mm broad, lanceolate. Inflorence terminal, with 3 cymes, 
bifurcate or ternately branched, rarely with one more cymes in axils of 
upper whorl of leaves, few-flowered. Pedicels somewhat longer than 
corolla, both glabrous. Corolla 3—7 mm across, infundibuliform, with 
short tube; lobes shorter 4 oblong, obtuse. Stamens on short filaments, 
equaling anthers; anthers exserted from throat of flower. Style short, 
included in tube of flower, bifid from middle; stigma globose. Fruit 
binate, mericarps globose, 3—4 mm long and broad, covered with dense, 
shiny hooked hairs on tubercles. Flowering April to June. Fruiting 
July to September. 

Coniferous and broad-leaved forests, especially ash, oak, beech, 
in deep shade on wet soil.—European USSR: All regions, besides 
Karelia-Lapland, and only in southernmost part of Dvina-Pechora. 
Caucasus: All regions. Western Siberia: Altai. Eastern Siberia: Angara- 
Sayan. Soviet Far East: Ussuri, Sakhalin (in Kuril Islands). Soviet 
Central Asia: Balkhash Region, Dzhungaria-Tarbagatai. General 
distribution: Scandinavia, Central Europe, Atlantic Europe, western 
and eastern Mediterranean, Balkans-Asia Minor, Iran-Afghanistan 
(northern part), Japan, China, North America (introduced). Described 
from Sweden, type in London. . 

Note. It should be noted that this species, insofar as its habitat is 
closely related with deep shade and wet humid soil, covers its wide 
range discontinuously, sometimes in extremely disjunct spots. The 
plant does not persist on loggings of more than a year. The size of the 
leaves and flowers and the type of pubescence on the leaves vary. In 
European and especially in Caucasian plants, the leaves are covered 
with appressed, short, scattered hairs on both sides, and in specimens 
from Asia, the leaves are predominantly glabrous, without setaceous 
hairs even along the vein beneath. But this is not a constant character. 

Economic importance. The whole plant, but particularly the 
flowers, contains coumarin in considerably larger quantities than other 
plants. In Western Europe, it has long been used for providing flavoring 
in Rhine wine and preparing “May wine.” In the Baltic Republics it 
was mixed, in the olden days, in the dry form with tobacco to render 
it fragrant (Maltsev in Tr. Byuro po prikl. bot. (1909) 585). 


Section 4. Cynancuica! DC. Prodr. IV (1830) 582, p.p.—Genus 
Cynanchica Fourr. in Ann. Soc. Linn. Lyon. nouv. sér. XVI (1868) 
398.—Inflorescence dichasial, of varied shapes, paniculate, spicate, 
almost capitate or sometimes pseudowhorled, with almost sessile 
flowers or on pedicels, often forming dense clusters at ends of branches, 


1Treatment by M.V. Klokov. 


219 


208 


surrounded by floral leavés. Margin of calyx often with less distinct, 
rarely with long teeth. Corolla infundibuliform or tubular- 
infundibuliform to nearly claviform, pink, lilac, light blue, whitish or 
yellowish, 4-merous with lobes shorter than tube or almost as long. 
Fruit rugose, covered with hemispherical or cuneate tubercles, rarely 
flat or scaly outgrowths, very rarely almost smooth. Subshrubs with 
woody, branched rootstock and numerous flowering stems, 4-sided, 
rarely almost terete at base. Leaves narrowly linear, more or less 
acuminate, always 4 in a whorl. 

In section Cynanchica DC. there are over 100 species, distributed 
widely around the entire Mediterranean Sea and in the adjoining 
countries to the north and east. This section, studied in detail on a 
worldwide scale by V.I. Kreczetowicz, represents a fairly natural group 
within the limits of the genus Asperula L. However, Kreczetowicz’s 
treatment, unfortunately, remained uncompleted and was not published 
after the untimely death of the author. Only fragments of this work 
have become known, which concern the flora of individual regions of 
the Soviet Union, published by the author in the collective work “Flora 
Yugo-Vostoka Evropeiskogo Rossii (Flora of southeastern European 
Russia) (1936) or in the first edition (1934) of “Flora Kavkaza” (Flora 
of the Caucasus) by A.A. Grossheim. The manuscript itself was long 
thought to be lost, but a considerable portion of it, containing the 
general system of classification of the subgenus and the Latin diagnoses 
of the species established or amended by the author, was recently 
discovered and has been fully incorporated in the present work, of 
course along with necessary corrections and additions. Kreczetowicz’s 
planned general system is given in still a preliminary form, with 
reservations regarding its content. The uniqueness of this system is in 
the aggregating of series into cycles, which is not done in Flora USSR, 
and in the naming of series according to the nature of the morphological 
characters and not according to the name of a typical representation of 
the series, as is done in the present work. On the other hand, the 
division of the sections into subsections, apparently needed here, but, 
requiring various preliminary studies for its basis, is not carried out. 
Within the limits of our flora, it is possible to denote two subsections; 
to the one belong the obligate psammophilous plants covered with 
scaly outgrowths, comparatively large fruits (the Lepidophorae cycle 
of Kreczetowicz’s system, with the best known representative being A. 
graveolens M.B.), and to the other belong all the remaining species 
(the Cynanchicae, Parviflorae, Racemiferae, Corymbiferae and 
Pontigenae cycles of his system), characteristic mainly of rocky 
outcrops, with small fruits covered with tiny tubercles. 


220 


209 


The studies conducted by Kreczetowicz convincingly established 
that this section can be differentiated into geographical races, adapted 
to specific, more or less limited territories, in terms of zones and 
regions, and to specific ecological conditions. In other words, in this 
section belongs the Mediterranean type of differentiation into narrow- 
area species (term of S.V. Yuzepchuk). Thus, the natural units of 
differentiation, which, according to the principles of V.A. Komarov’s 
school, can qualify as species, are here geographical races; the latter 
in this case are quite easily distinguishable by a complex of 
morphological characters. The uniting of these species— geographical 
races—into natural groupings of series presents considerably greater 
difficulties, which Kreczetowicz had also encountered. The clear pattern 
of species differentiation, in this case, is completely explained by the 
absence of free interspecific hybridization. Such hybridization could 
obscure basically the same pattern, for example, in the genus Thymus. 
But the difficulty of establishing series or higher natural groups can 
be explained by the complex and less understood phylogenetic 
interrelationship between the species and species groups within the 
section Cynanchica (Fourr.) V. Krecz. 

After Kreczetowicz, the only paper devoted to species of this 
section was that of M.G. Popov and V.G. Chrshanovsky under the 
title: Sistematicheskii analiz vidov, ob’edinyaemykh pod nazvaniem 
Asperula cynanchica na Ukraini. K voprosu ob endemizatsii ukrainskoi 
flory (Taxonomic Analysis of the Species Combined under the Name 
Asperula cynanchica in the Ukraine. On endemism in the Ukrainian 
flora) (1945). In this paper the authors approach the problem from 
their special viewpoint, while clearly underestimating the difficulty. 
Examination of the Ukrainian species of the section proposed by the 
authors of this system showed that they, on the one hand, incorrectly 
distinguished the species A. pseudograveolens M. Pop. et Chrshan. 
and A. maeotica M. Pop et Chrshan., which do not represent separate 
geographical races and are not distinguished from the earlier described 
A. graveolens M.B. (first species) and A. stevenii V. Krecz. (the second 
one), and, on the other hand, incorrectly combined into a single species 
(A. tephrocarpa Czern. ex M. Pop. et Chrshan.) several independent 
geographical races belonging to different series and cycles of the system 
proposed by Kreczetowicz. Moreover, they established the concept of 
a “conglomerate, probably, of hybrid forms between A. cynanchica 
and the present A. setulosa,” despite the fact that these species cannot 
hybridize and are never encountered at the same place together. The 
attempts made in this paper to use for a floristic taxonomy the 
elementary morphological “Linnaeon” [broad] and “Jordanon” [narrow] 
concepts modernized on a hybrid-genetic basis (this approach was 


223 


210 


used again and again by M.G. Popov in his various treatments) did not 
produce positive results here. Evidently, use of the indicated terms 
[Linnaeon and Jordanon] did no more than confuse the matter further. 
The problem of natural groupings of geographic races cannot be 
resolved on a formal morphological base, by way of arbitrarily 
established ‘Linnaeons’; it can be resolved only by the application of 
the historical method, using phylogenetic analysis and, consequently, 
(phylogenetic) synthesis. 


Series 1. Eucynanchicae V. Krecz.—Almost entirely herbaceous 
plants or more typically subshrubs, forming mats. Cauline leaves 
narrow, linear to almost subulate, flat or with recurved margin, floral 
leaves mostly oblong-lanceolate, much smaller, shorter than flowers 
and carpels. Inflorescence paniculate. Flowers on distinct short pedicels. 
Corolla whitish, infundibuliform or tubular-infundibuliform, smaller 
moderately large (2-2.5—4.5 mm long); lobes oblong-lanceolate, 
mucronate, almost equaling tube, densely setulose with less distinctly 
thickened tip. Style 0.5-2 mm long free at tip, with capitate stigmas. 
Fruit very small, 1.5-2.25 mm long finely tuberculate. 

A central European and partly eastern European series, to which, 
besides our species, several western European mountain species belong. 
The members of this series are distinguished from the rest of the 
subgenus as the most mesophilic. The greatest xeromorphism among 
them can be found in the specialized species of calcareous outcrops 
(A. tephrocarpa Czern. and A. creticola Klok.), to a certain extent 
transitional toward the following series. According to the Kreczetowicz 
system, this series belongs to the cycle Cynanchicae V. Krecz. along 
with the western European series Tenuiflorae V. Krecz. 


13. A. semiamicta Klok. sp. n. in Addenda XXII, 694. 

Perennial. Poorly expressed herbaceous green subshrub. Flowering 
stems fairly numerous, ascending, 30-45 cm tall, densely setulose in 
lower part together with leaves, glabrous and smooth only above the 
middle, like the upper leaves. Lower leaves elliptical to obovate and 
oblong-obovate, 2.5—-5 mm long, 1.25—2 m broad, narrowed into petiole 
at base, mucronate, middle cauline leaves from linear-oblong (lower) 
to narrowly linear 10-43 mm long, 1.25-1.5 mm broad, cuspidate, 
upper leaves (in inflorescence) linear or oblong-lanceolate, 2.5— 8 mm 
long, 0.6—1 mm broad, acuminate, glabrous, entire; floral leaves oblong 
or ovate-lanceolate, about 2 mm long, 0.6-0.75 mm broad, finely 
acuminate, smooth. Inflorescence narrowly paniculate with numerous, 
moderately long or short branches. Flowers on short, often less distinct 
pedicels. Corolla infundibuliform, 2.25—2.75 cm long; lobes oblong- 


211 


Bee 


Plate XII. 

221 1. Asperula stevenii V. Krecz., general appearance of plant, portion of 
inflorescence, flower.—2. A. tephrocarpa Czern., general appearance of plant, 
portion of inflorescence, flower.—3. A. cynanchica L., general appearance of 

plant, portion of inflorescence, flower. 


224 


212 


lanceolate, mucronate about 1.25 mm long and 0.5 mm broad, whitish, 
densely setulose. Style up to 1.5 mm long. Anthers about 1 mm long. 
Fruit 1.5-2 mm long, finely tuberculate. June to August. 

Stony slopes.— European USSR: Upper Dniester, Middle Dnieper 
(western part). Endemic. Described from vicinity of Zaleshchiky on 
Dniester. Type in Leningrad, cotype in Kiev. 

Note. This species is known from several localities along the 
Dniester and its tributaries. Our observations show that it is abundant 
there. In all probability, it is one of a number of endemic plants of the 
Dniester Region. It is extremely close to true A. cynanchica L. s. str., 
but differs sharply from it in morphological features, which are more 
archaic in A. semiamicta Klok. 


14. A. cynanchica L. Sp. pl. (1753) 104; Lbd. Fl. Ross. II, 1 
(1844), excl. var. var. p.p.; Schmalh. Fl. IT, 9, excl. var. var. p.p.; 
Chrshan. in Vizn. roslin URSR, 309 p.p.— Exs.: Fl. exs. austro-hung. 
No. 2230; Fl. Hung. exs. No. 687. 

Perennial. Almost entirely herbaceous, bright green plant. 
Flowering stems several, rarely numerous, suberect or ascending from 
base, 15-60 cm tall, fairly slender, occasionally somewhat curved, 
weakly scabrous-setulose or scabrous in lower part, glabrous and 
smooth above, divaricately branched from middle or below, rarely 
above, comparatively densely foliated. Lower leaves from ovate- 
semicircular to oblong-elliptical or oblong-obovate, 0.75—6 mm long, 
0.5—2 mm broad, narrowed at base into a petiole: at base middle cauline 
leaves narrowly linear, 9-50 mm long, 0.75—2 mm broad, often up to 
30-35 mm long and 1.25—1.5 mm broad, flat or nearly so (with slightly 
revolute margin), somewhat narrowed toward base; lower leaves weakly 
scabrous along margin, the rest smooth; floral leaves linear or oblong- 
lanceolate, 1.25-1.5 mm long, 0.3-0.75 mm broad. Inflorescence 
paniculate. Flowers on extremely short or more distinct up to 2-3 mm 
long pedicels. Corolla tubular-infundibuliform, 3—4.5 mm long; lobes 
oblong-lanceolate, mucronate, 1.5—2.25 mm long, above 0.6 mm broad, 
white or very pale pink, scabrous with very fine less distinct hairs. 
Styles 1-2 mm long, free at top or almost from base. Anthers 
lanceolate, about 1 mm long. Fruit 2-2.5 mm long, densely covered 
by fine vesicular tubercles. June to August (Plate XII, fig. 3). 

Steppe slopes, meadows steppe, dry meadows, open forests along 
fields and forest clearings.— European USSR: Upper Dniester, Upper 
Dnieper (southern part), Upper Volga (in extreme south), Bessarabia 
(northern part), Middle Dnieper, Volga-Don, Black Sea Region 
(northern part), Lower-Don (northern part). General distribution: 


225 


213 


Central Europe (Austria, Hungary, Poland). Described from Central 
Europe. Type in London. 

Note. A highly variable plant, showing differences with changing 
habitat conditions. Probably, it hybridizes with closely related species, 
although in this section hybrids or hybridogenic forms have not yet 
been confirmed with certainty. As accepted here, A. cynanchica cannot 
be regarded as a collective species [species group]. Kreczetowicz 
distinguished A. cynachica L. s. str. from A. michelsonii V. Krecz., 
named by him in the herbarium, although a description of the latter is 
missing from his manuscripts. A brief diagnosis of this still problematic 
species was published by the present author in ‘Viznachnik roslin 
URSR’ (Manual of the plants of the Ukr. SSR) (1950, p. 309). This 
species is well-distinguished from A. cynanchica L. s. str. in its habit, 
since it forms a dense, pulvinate mat 8-12 mm tall; the middle cauline 
leaves are 15-10 mm long; the stems are densely leafy; the corolla is 
small, with the lobes longer than the tube, whitish, violet inside; the 
fruit is about 2 mm long, with subacute tubercles. The type comes 
from the vicinity of Zbarazh, Ternopol Region of UkrSSR, where the 
plant was collected on limestone hills by A.I. Mikhelson. Unfortunately, 
this unique form does not reappear in the new collections; the authentic 
material is insufficient to prove convincingly that it represents a distinct 
geographic race. It should be noted that plants from the eastern regions 
of the range indicated here for A. cynanchica L. (e.g. distributed in 
the Herbarium of the Russian Flora as var. elongata Stev.) differ 
considerably from the typical Central European race in their narrower 
leaves and, in all probability, ger to the independent Eastern 
European race. 


15. A. creticola Klok. sp. n. in Addenda XII, 694.—A. cynanchica 
auct. pro min. parte.—Exs.: GRF, No. 1668 (sub. A. cynanchica var. 
elongata Stev.).' 

Perennial. Grayish green subshrub, forming lax mat. Flowering 
stems numerous, ascending, 13-30 cm tall, more or less distinctly 
scabrous-setulose in lower part, glabrous and smooth above middle, 
branched from middle or below. Basal leaves from ovate-deltoid to 
oblong, 0.5—4 mm long, 0.75-1 mm broad, more or less scabrous; 
middle cauline leaves linear-subulate, 7-26 mm long, 0.25-0.75 mm 
broad, acuminate, or cuspidate with revolute margin, those situated at 
bottom scabrous the rest smooth; floral leaves oblong- or linear- 
lanceolate, 2-2.5 mm long, 0.5-—0.75 mm broad. Inflorescence 


'A part of the distributed specimens; the other part belongs to A. cynanchica L. 


226 


214 


paniculate, with divaricate branches. Flowers on short, but more or 
less distinct pedicels. Corolla infundibuliform, 2.5—3 mm long; lobes 
reflexed, oblong-lanceolate, 1.25—1.75 mm long, whitish, scabrous- 
setulose. Fruit about 1.75 mm long, finely tuberculate. June to August. 

On bare chalks and marls and steppe slopes with calcareous subsoil, 
apparently, also on limestones.— European USSR: Middle Dnieper 
(southeastern part), Volga-Don (southwestern part), Black Sea Region 
(eastern part), Lower Don (northwestern part). Endemic. Described 
from Svyatye Gory along northern Donets. Type in Kiev. 

Note. This morphologically well-defined species, which is clearly 
distinguished from both A. cynanchica L. and A. tephrocarpa Czern., 
occupies a middle position between them in this series. It often occurs 
together with A. tephrocarpa Czern. on calcareous outcrops, occupying, 
however, a somewhat lower position. The origin of the species is 
unclear. Possibly, it is a hybrid form, though by no means a recent 
hybrid between the named species. It should be noted that, according 
to our observations, A. creticola Klok. often occurs on calcareous 
outcrops along the Krynka River in the Azov Sea area, where both A. 
exasperata Czern. and A. cynanchica L. are absent. 


16. A. tephrocarpa Czern. in herb. ex M. Pop. et Chrshan. in 
Byull. Mosk. Obshch. ispyt. Prip. otd. biol. L. 5-6 (1945) 96, p.p. 
emend. Klok.—A. tephrocarpa var. densa M. Pop. et Chrshan. l.c. 
97.—A. supina auct. fl. ucr. nonnul. non M.B.—A. exasperata auct. 
fl. ucr. nonnul. non V. Krecz. sensu proprio. 

Perennial. Grayish green subshrub, forming dense, often pulvinate 
mats. Flowering stems numerous, ascending, 3—20 cm tall, not very 
slender, more or less distinctly scabrous-setulose throughout, branched 
in upper part, densely foliated. Basal leaves from ovate-deltoid, obtuse 
to oblong, shortly mucronate, 1-3 mm long, 0.5—1 mm broad, scabrous- 
setulose, middle cauline leaves linear-subulate, 5-17 mm long, 0.25-— 
0.75 mm broad, short-acuminate, with revolute margin more or less 
scabrous; upper leaves (in inflorescence) oblong-lanceolate, up to 4 
mm long, and 1.1 mm broad, scabrous; floral leaves oblong or linear- 
lanceolate, 1-3 mm long, 0.25—0.75 mm broad, cuspidate, scabrous. 
Inflorescence shortly paniculate. Flowers on very short, often barely 
distinct pedicels, more or less crowded at ends of branches of 
inflorescence. Corolla infundibuliform, 3—3.75 mm long, with oblong- 
lanceolate, mucronate, 1.5—-2 mm long, 0.6—0.8 mm broad, sharply 
deflexed; lobes, whitish, scabrous-setulose. Styles 0.5-1 mm long, 
free above the middle. Anthers oblong, about 1 mm long. Pistil 1.75- 
2 mm long dark brown, densely covered with whitish vesicular 
tubercles. Mericarps highly curved. May to August (Plate XII, fig. 2). 


227 


215 


On calcareous outcrops.— European USSR: Middle Dnieper 
(southeastern part), Volga-Don (southwestern part), Lower Don 
(northern part). Endemic. Described from the chalks along Seversky | 
Donets River. Type in Kiev. 

Note. This species, characteristic of calcareous outcrops of the 
Donets Basin, was well distinguished from all others by Prof. V.M. 
Czernajew; which is fully documented by the material in his herbarium. 
Kreczetowicz referred it to his A. exasperata V. Krecz., which 
undoubtedly is an error. M.G. Popov and V.G. Chrshanovsky, in 
restoring Czernajew’s species, gave it an unnaturally broad 
circumscription. They also referred to A. tephrocarpa Czern. some 
species related to A. rumelica Boiss., although A. tephrocarpa Czern. 
differs from them even more than from true A. cynanchica L.; the 
typical form of the species [A. tephrocarpa], growing on limestone, 
was recognized as a separate “variety’—var. densa M. Pop. and 
Chrshan. A. tephrocarpa Czern. sensu proprio and A. creticola Klok. 
in many respects differ from A. cynanchica L. and the other Central 
European species of the series, forming as it were, a transition from 
the Eucynanchicae to the eastern series Petraeae, probably derived 
from the former. These two species evidently should be combined into 
a separate intermediate series Tephrocarpae Klok. 


Series 2. Petraeae Klok. Small subshrubs, forming mats. Cauline 
leaves linear-subulate, with recurved margin. Inflorescence paniculate. 
Flowers often almost sessile, rarely on distinct but extremely short 
pedicels. Corolla very small (2.25-3 mm long), with oblong-ovate- 
lanceolate, subobtuse lobes, whitish, glabrous and almost smooth. Style 
extremely short, not longer than 1 mm. Fruit 2-3 mm long, very 
finely tuberculate. Similar to preceding series in other features. 

An eastern, in relation to the preceding series, Volga-Ural 
petrophilous series. It clearly differs from the previous series 
Eucynanchicae V. Krecz. by having a glabrous corolla and 
comparatively large fruits. Kreczetowicz referred both known species 
of this series to his series Leianthae V. Krecz., the type of which is 
A. leiantha A. Kern.—an endemic species of the southern Tirol having 
flowers with much larger, yellowish pink corollas, clearly having no 
immediate relationship to our Volga-Ural species. We here segregate 
the Volga-Ural Leianthae into an independent series, which will be a 
more realistic approach. 


17. A. exasperata V. Krecz. in Fl. Yugo.-Vost. V1 (1936) 256 
(rossice); in Addenda XXII, 695. 


228 


216 


Perennial. Dark green subshrub with weakly developed sterile 
shoots, forming divaricate or flattened mats. Flowering stems suberect 
or ascending, 5—25 cm tall, not very slender, scabrous with cuneate 
papillae to the very top, distinctly so at nodes, branched from above 
or below middle, comparatively densely foliated. Basal leaves deltoid, 
ovate, oblong-ovate or oblong, 1-4 mm long, 1—1.5 mm broad, weakly 
scabrous, middle cauline leaves from linear-spatulate, distinctly 
broadened toward tip, to linear-subulate (upper ones), 6-15 mm long, 
0.5—1 mm broad, mucronate, flattened, thickened along margin, rarely 
with somewhat revolute margin, fairly weakly scabrous; upper leaves 
(in inflorescence) oblong-lanceolate, about 3 mm long and 1 mm broad 
and weakly scabrous; floral leaves oblong or oblong-ovate-lanceolate, 
1.75-2 mm long, 0.75—1 mm broad, acuminate, weakly scabrous to 
almost smooth. Inflorescence paniculate, more or less divaricate or 
condensed. Flowers subsessile or rarely on distinct, up to 1 mm long, 
pedicels, (2)3 together at ends of short branches of inflorescence. 
Corolla fairly broadly infundibuliform, 2.25—2.75 mm long; lobes more 
or less deflexed, oblong-ovate-lanceolate, obtuse, 1.25—1.5 mm long, 
0.4-0.5 mm broad, whitish, with tapering thickened tip; corolla tube 
smooth or almost smooth, lobes with scattered, very short, but 
comparatively broad setae along margin. Style 0.4—-0.7 mm long. 
Anthers about 1 mm long. Ovary pointedly tuberculate. Fruit 2-3 mm 
long, wrinkled and finely tuberculate, yellowish brown. June to August. 

Calcareous outcrops.— European USSR: Volga-Don (eastern part), 
Lower Don (northeastern part). Endemic. Described from vicinity of 
Khvalynsk-on-Volga. Type in Leningrad. 

Note. This species is rather common in the Volga calcareous 
regions (Khvalynsk, Volsck, Belaya Glinka and other places), and it 
also occurs on calcareous outcrops in the Don Basin along the Khoper 
River, and probably also along the Medveditsa. Inasmuch as the species 
was described by Kreczetowicz from the Volga calcareous region, it 
became necessary for us to select as the type another specimen, fitting 
the description better, instead of the Ukrainian specimen provisionally 
regarded by the author as the type, which belongs to A. tephrocarpa 
Czern. The necessity of such a change was agreed to by the author of 
the species some time ago. A. exasperata V. Krecz. and A. tephrocarpa 
Czern. acutally are very close to one another in external features, a 
fact essentially explained by the identical action of the calcareous 
substrata on them; however, they sharply differ from one another 
by the characters of the series, which are more profound but not so 
evident. 


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217 


18. A. petraea V. Krecz. in Fl. Yugo-Vost. VI (1936) 254 (rossice); 
in Addenda XXII, 696.—A. cynanchica Korsh. Tent. (1898) 189, non 
L.—Ic.: Fl. Yugo-Vost. VI, fig. 649. 

Perennial. Bright green, lustrous subshrub, with well-developed 
sterile shoots, forming loose mats. Flowering stems obliquely 
ascending, 7-28 cm tall, slender, weakly scabrous-setulose in lower 
part, glabrous and smooth in upper part, more or less branched above 
the middle. Basal leaves obovate, oblong-obovate or spatulate, 1-4 
mm long, 0.5—1 mm broad, narrowed at apex; middle cauline leaves 
narrowly linear, 6-21 mm long, 0.5-0.75 mm broad, mucronate or 
cuspidate, with more or less revolute margin, extremely mildly scabrous 
upper leaves (in inflorescence) oblong-lanceolate, 2-3 mm long, 0.5— 
0.7 mm broad, acuminate, smooth; floral leaves of same shape or 
lower relatively broader, oblong-ovate-lanceolate, commonly about 1.5 
mm long, and 0.5 mm broad. Inflorescence paniculate, fairly lax, but 
flowers almost sessile, in small 3-flowered clusters at ends of more or 
less elongated branches. Corolla infundibuliform, about 3 mm long; 
lobes slightly deflexed, oblong-ovate-lanceolate, mucronate, about 1.5 
mm long, 0.5-0.7 mm broad, whitish, glabrous and smooth. Style 
about 1 mm long; fruit 3—4 mm long, yellowish brown, slightly rugose 
covered with very fine whitish tubercles. June to August. 

Limestone-stony slopes and outcrops.— European USSR: Volga- 
Don (Zhiguli), Volga Region, Volga-Kama (southern part). Western 
Siberia: Upper Tobol (southwestern part). Soviet Central Asia: Aralo- 
Caspian (northwestern part). Endemic. Described from northern 
extremity of Southern Urals (Mt. Egazinskaya near Kyshtym in 
Chelyabinsk Region). Type in Leningrad. 


Series 3. Supinae Klok.—Small subshrubs with well-developed 
sterile shoots, forming dense mats. Cauline leaves narrowly linear, 
fairly short, flattened; floral leaves comparatively large, exceeding 
carpels, often ovate-lanceolate, with dentate margins. Inflorescence 
corymbose. Flowers almost sessile, compacted at ends of few branches 
of inflorescence in dense, capitate clusters surrounded by floral leaves. 
Corolla very small or larger (2-5 mm long), pink, with setulose or 
glabrous and almost smooth tube; lobes smooth oblong-ovate- 
lanceolate, obtuse, approximately as long as tube. Style short (0.5-1 
mm long). Fruit 1.5-2 mm long, finely tuberculate. 

To this series we refer three Crimean mountain species. 
Kreczetowicz, evidently, did not differentiate them and referred A. 
supina M.B. s.l. to series Capitellatae V. Krecz., the type of which is 
the Asia Minor species A. capitellata Hausskn. et Bornm., probably 
not so close to the endemic Crimean species of our series. This series, 


230 


218 


as also both of the preceding ones, is characterized by elongated corolla 
lobes, approximately as long as the tube. In all remaining series of our 
flora (not considering the isolated series Graveolentes Klok., 
corresponding to the cycle Lepidophorae V. Krecz.) the corolla lobes 
are distinctly or considerably shorter than the tube. We consider 
probable the phylogenetic closeness of the first three series, although 
according to the Kreczetowicz system they belong to different and not 
closely related cycles. 


19. A. supina M.B. FI. taur.-cauc. I (1808) 101 and III (1819) 
103; DC. Prodr. IV, 383, quoad plantam tauricam.—A. cynanchica B. 
supina (M.B.) Schmalh. Fl. II (1897) 9, p.p. 

Perennial. Subshrubs with well developed sterile shoots, forming 
fairly dense mats, dark-green in young parts and yellowish-greenish 
in older parts. Flowering stems suberect or obliquely ascending 
(reclining), about 10-12 mm tall, with up to 1-1.5 mm long internodes, 
not slender (about 1 mm thick), scarbous-setulose throughout, sparsely 
so upwards, bifurcately or ternately branched from middle or below, 
branches directed obliquely upward, comparatively densely foliated. 
Basal leaves oblong-lanceolate, obtuse, very small, poorly preserved; 
cauline leaves narrowly linear to narrowly linear-lanceolate upwards, 
3-12 mm long, 0.5—-1 mm broad (at base) cuspidate, stiff, with 
prominent thick veins on lower side, and extremely narrow bands of 
tissue along sides, scabrous-setulose along margin, more or less 
deflexed from stems, but almost straight; floral leaves oblong-ovate- 
lanceolate, 2-3 mm long, 0.75—1.25 mm broad, narrowly acuminate at 
apex, irregularly serrulate along margin in lower part. Branches of 
inflorescence somewhat divaricate, flowers sessile or subsessile, at 
end of last branches of inflorescence in a dense 5—9-flowered capitate 
cluster, surrounded by floral leaves. Corolla infundibuliform, 2.5—3 
mm long; lobes deflexed, oblong-ovate-lanceolate, obtuse, 1.25—1.5 
mm long, about 0.5 mm broad, pale pink; tube comparatively broad, 
densely setulose; lobes glabrous and smooth, with less distinct 
thickened tip. Style about 0.5 mm long. Anthers oblong, about 1 mm 
long. Fruit 1.5—2 mm long, finely tuberculate. June to August (Plate 
XIII, fig. 2). 

Calcareous mountain slopes.— European USSR: Crimea (known 
only from the calcareous mountains in vicinity of Belogorsk). Endemic. 
Described from the indicated place. Type in Leningrad. 

Note. In the treatment of this species we follow S.V. Juzepczuk, 
who restored the correct concept when establishing his new species A. 
caespitans Juz. 


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219 


20. A. caespitans Juz. in Bot. mat. Gerb. Bot. inst. AN SSSR... 
XIV (1951) 35.—A. supina auct. fl. taur. pro. max. parte, non M.B.— 
Ic.: Juzepcezuk, l.c. fig. 3. 

Perennial. Dark green subshrub, with numerous densely foliated, 
sterile shoots. Flowering stem from decumbent base ascending 
obliquely or suberect, S—15(25) cm tall, slender (0.4—0.7 mm thick), 
with up to 0.8-3 cm long internodes, puberulent throughout or 
sometimes only below nodes, fairly weakly furcately or ternately 
branched above. Basal leaves from short oblong, obtuse, to oblong- 
lanceolate, mucronate, 1-3 mm long, 0.5—-1 mm broad, considerably 
longer than corresponding, greatly shortened internodes, margin smooth; 
cauline leaves narrowly linear to narrowly linear-lanceolate upward, 
3-11 mm long, 0.5—1 mm broad (at base), vein thickened underneath 
and extremely narrow tissue bands on its sides, scabrous-setulose along 
margin or, puberulent as in stem, sometimes also pubescent below 
along vein; floral leaves oblong or ovate-lanceolate, 2-4.5 mm long, 
0.75-2 mm broad, narrowly acuminate, more or less crenate and 
glabrous or sometimes ciliolate along margin. Inflorescence compact. 
Flowers sessile or subsessile, at end of branches of the first order in 
dense, often almost capitate 3—9-flowered clusters, not more than 3 | 
such clusters in inflorescence, normally, lying very close at end of the 
stem, forming an overall lax cluster. Corolla tubular-infundibuliform, 
3.5-5 mm long; lobes oblong-ovate or oblong-ovate-lanceolate, 
mucronate, 1.5—2.5 mm long, 0.75—1 mm broad, more or less reflexed, 
pink, with whitish, thickened tip; corolla tube densely or diffusely 
setulose, lobes glabrous along margin. Style 0.5—1 mm long, free at 
tip. Anthers about 1 mm long. Fruit about 2 mm long covered fine, 
obtuse tubercules. June to August. 

Mountain slopes.— European USSR: Crimea (mainly in Yaila). 
Endemic. Described from Nikitskaya Yaila. Type in Leningrad. 


21. A. tranzshelii Klok. sp. n. in Addenda XXII, 697.—A. supina 
auct. Fl. taur. pro min. parte. 

Perennial. Yellowish green subshrub, with numerous, densely 
foliated, sterile shoots, forming fairly dense mats. Flowering stems 
from procumbent or ascending base to suberect, 10-20 cm tall, with 
up to 2-4 cm long internodes, not very slender (about 1 mm thick), 
glabrous and smooth or sometimes scabrous only at base weakly, 
furcately or ternately branched above. Basal leaves linear- or oblong- 
lanceolate, 2-3.5 mm long, 0.5—1 mm broad, obtuse or more or less 
acuminate; cauline leaves narrowly linear to narrowly linear-lanceolate, 
4-14 mm long, 0.5-1 mm broad, cuspidate or flattened or with 
somewhat revolute margin, weakly scabrous-setulose along margin 


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(except for the upper ones which are smooth); floral leaves oblong- 
ovate-lanceolate, 2-4 mm long, 0.75-1.5 mm broad, narrowly 
acuminate, irregularly crenulate along margin. Inflorescence compact. © 
Flowers almost sessile or on very short (about 0.5 mm long), less 
distinct pedicels, at ends of first order branches in compact, almost 
capitate 3—9-flowered clusters, or more rarely in several less dense 
clusters at ends of branches of second order. Corolla infundibuliform, 
about 4.5 mm long, lobes reflexed, oblong-ovate-lanceolate (ligulate), 
subobtuse or obtuse about 2.5 mm long and 1-1.25 mm broad, pale 
pink, with whitish, wrinkled tip; corolla tube fairly broad, weakly 
scabrous from cuneate papillae or almost smooth; lobes smooih. Styles 
0.5—1 mm long, free from base. Anthers oblong, 1-1.25 mm long. 
Ovary finely tuberculate. Fruit unknown. June to August. 

Mountain slopes.— European USSR: Crimea (southern mountain 
slopes). Endemic. Described from environs of Yalta. Type in Leningrad. 


Series 4. Stevenianae Klok.—Flowering stems comparatively tall 
(25-65 cm), velutinous in lower part together with leaves. Middle 
cauline leaves linear-subulate, elongated (up to 35-45 mm long), with 
revolute margin; floral leaves oblong- or linear-lanceolate, very small. 
Inflorescence paniculate. Flowers on distinct pedicels. Corolla yellowish 
pale pink, tubular-infundibuliform, large, 4.5—-8 mm long, with 
elongated (not shorter than 3 mm) tube and short (2-4 times as short 
as tube), obtuse or subobtuse lobes, scabrous-setulose. Style 2—3.5 
mm long. Fruit up to 2.25 mm long, with tiny, obtuse tubercles. 

Crimea-New Russian species, which also occur along the northern 
coasts of the Black Sea and Sea of Azov. 

Opinions differ on the origin of the basic species of this series A. 
stevenii V. Krecz. Kreczetowicz refers it to the eastern Mediterranean 
cycle Racemiferae V. Krecz, and places it in the same series 
(Gracilentae V. Krecz.) with the Banatian species A. tenella Heuff. 
M.G. Popov et V.G. Chrshanovsky refer it (under the name A. maeotica 
M. Pop. and Chrshan.) also to the eastern Mediterranean group 
Longiflorae auct. and place it close to A. aristata L. fil. (1.c.). The 
eastern Mediterranean connection here is quite obvious, but also clear 
is the presence of a noticeable gap in affinities, which allows us to 
segregate A. stevenii V. Krecz. and the species closest to it into a 
separate southern Pontic series. 


22. A. stevenii V. Krecz. in Grossh. Fl. Kavk. IV (1934) 125; 
Klokov in Vizn. roslin URSR, 309; juzepczuk. in Bot. mat. Gerb. Bot. 
inst. An SSSR, XIV, 34.—A. cynanchica 8. elongata Stev. in DC. 
Prodr. IV (1830) 583.—A. cynanchica M.B. FI. taur.-cauc. I (1808) 


233 


D2A 


100, quoad pl. taur. pro max. parte, non L.—A. longiflora C. Koch in 
Linnaea, XXIII (1850) 460, non Waldst. and Kit.—A. maeotica M. 
Pop. et Chrshan. in Byull. Mosk. obshch. ispyt. prir. L, 5-6 (1945) 
97-98. 

Perennial. Grayish green subshrub, with shortened, woody branches 
of main axis, sterile shoots not developed. Flowering stems ascending 
at base or almost erect, 25-65 cm tall, obtusely 4-sided, comparatively 
slender, densely velutinous with short, about 0.25 mm long fairly sort 
hairs in lower part, glabrous and smooth or nearly so above middle, 
furcately or ternately branched above, with numerous, moderately long 
branches, comparatively sparsely foliated. Basal leaves from ovate- 
semiorbicular to elliptical, distinctly narrowed at base, 1-3 mm long, 
about 1 mm broad, pubescent; middle cauline leaves linear-subulate, 
8-35 mm long, 0.5—1.5 mm broad, short-acuminate, with more or less 
revolute margin, just like the stem those situated below pubescent, the 
remaining glabrous; upper (in inflorescence) leaves linear- or oblong- 
lanceolate, 2-5 mm long, 0.3—0.75 mm broad, more or less narrowly 
acuminate, glabrous and smooth (and also along margin); floral leaves 
similar in shape or somewhat broader at base, 1.5—2.25 mm long, 0.3-— 
0.4 mm broad, smooth margin. Inflorescence paniculate, not broad. 
Flowers on short, 0.5—1.5 mm long, thicker pedicels, situated at ends 
of short branches of inflorescence, (1)2—3 together. Corolla tubular- 
infundibuliform 5.5-7(8) mm long; lobes weakly recurved, oblong- 
elliptical or ovate, obtuse, 1-2 mm long, 0.5—0.75 mm broad, yellowish 
pale pink; tube of corolla narrowly cuneate, elongated, not shorter 
than 3.5 mm, densely covered with cuneate, acute papillae, and thus 
highly scabrous; lobes also scabrous, on midrib and along margin, but 
somewhat weaker, without distinct thickened tip. Styles 3-3.5 mm 
long, free above middle, unequal. Anthers about 1 mm long. Fruit 
1.75-2.25 mm long, rust-brown, densely covered with fine, obtuse 
tubercles. June to August (Plate XII, fig. 1). 

On coastal shell-rock sands, also stony steppe or mountain 
slopes.—European USSR: Black Sea Region (Biryuchy Island in Sea 
of Azov), Lower Don (along coast of Sea of Azov), Crimea. Caucasus: 
Ciscaucasia (northwestern). Endemic. 

Note. The plants from Biryuchy Island do not differ from typical 
Crimean ones. Their description as the distinct species A. maeotica M. 
Pop. et Chrshan., which for some reason was not even compared by 
the authors with A. stevenii V. Krecz., is without foundation. The 
forms related to this species and restricted to the Stavropol uplands, 
probably representing a separate geographic race with less pubescence 
and smaller corollas, are in need of additional studies. Somewhat 
different also is the highe-mountain form of the species, collected in 


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the Yaila, although its differences, apparently, do not exceed the range 
of modifications of the plant. A form unique to the northern Black Sea 
coast is segregated by us as a distinct species and described below. 
Thus, A. stevenii V. Krecz. appears to be a southern Pontic steppe 
species with the typical Azov (circummeiotic) geographical range. 


23. A. bidentata Klok. sp. n. in Addenda XXII, 697.—A. flaccida 
Velen. Fl. Bulgar. (1891) 238 ex min. parte.—A. montana Grecescu, 
Consp. fl. Rom. (1898) 265, quoad pl. Dobrudsh.—A. stevenii V. Krecz. 
l.c. pro min. parte. 

Perennial. Subshrub with shortened, woody branches of the main 
axis and weakly developed sterile shoots. Flowering stems (up to 8 in 
number) suberect, 30—60 cm tall, velutinous with short hairs in lower 
part together with leaves, more or less branched above middle. Basal 
leaves from semiorbicular to oblong-ovate and obtuse or oblong- 
lanceolate and acuminate, 1.5—7 mm long, 1.25—1.5 mm broad, middle 
cauline leaves linear subulate, 20-43 mm long, 0.5—0.75 mm broad, 
thicker, short-acuminate, with closely revolute margin; floral leaves 
oblong- or linear-lanceolate, 1.5-2 mm long, 0.3-0.5 mm broad, 
acuminate, flattened, along margin glabrous and smooth. Inflorescence 
paniculate, not broad. Flowers on very short pedicels, 3-5 together, at 
ends of short branches of inflorescence, often, though not in all flowers, 
calyx with 2 narrowly linear-filiform 0.5-0.75 mm long setulose teeth. 
Corolla tubular-infundibuliform, about 4.5-4.75 mm long; lobes weakly 
deflexed, oblong-lanceolate or ovate-lanceolate, obtuse, about 1.5 mm 
long and 0.6-0.75 mm broad, whitish or very pale yellowish pink; 
corolla tube 3—3.5 mm long, setulose along with lobes; lobes with less 
distinct thickened tip. Styles 2-2.5 mm long, free at tip, unequal. 
Anthers about 1 mm long. Fruit about 2 mm long, finely tuberculate. 
June to August. 

On coastal, shell-rock sands.—European USSR: Bessarabia 
(southern part). General distribution: Balkans (Dobrudzha). Described 
from Tendra Sand Bar on the Black Sea. Type in Kiev. 

Note. Kreczetowicz did not distinguish this species from A. stevenii 
V. Krecz. distributed in the more eastern and southern regions, to 
which it is extremely close. Popov and Chrshanovsky (l.c.) referred 
specimens of it to their hypothetical “conglomerate of putative hybrid 
forms between A. cynanchica and A. setulosa.” However, it is not 
directly related to either of these species. 


Series 5. Rumelicae Klok.— Generally, weakly stiffened subshrubs 
without sterile shoots. Flowering stems mostly comparatively tall (up 
to 50 cm), weakly scabrous-setulose towards base, highly branched, 


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223 


often with very slender branches. Middle cauline leaves narrowly linear- 
subulate to almost filiform, elongated (up to 46 mm), with more or 
less revolute margin; floral leaves linear-lanceolate, very small (up to 
2 mm long, and 0.5 mm broad). Inflorescence paniculate. Flowers on 
extremely short (up to 0.5 mm long), rarely more distinct pedicels. 
Corolla usually very small, rarely larger (2—4.5 mm long) with obtuse 
lobes, one and half times as short as tube, whitish, scabrous-setulate. 
Styles 0.4-1.75 mm long. Anthers up to 2.5 mm long, finely 
tuberculate. 

A southern steppe Pontic species, not extending, however, beyond 
Ciscaucasia. The main species of the series, A. rumelica Boiss., was 
placed by Kreczetowicz in the cycle Parviflorae V. Krecz. and the 
series Leptocladae V. Krecz., placed in his system directly after the 
Encynanchicae V. Krecz. The species A. hypanica Klok., described by 
us, more closely connects this species with the former, and thus they 
may be regarded as closely related groups. 


24. A. hypanica Klok. sp. n. in Addenda XXII, 698. 

Perennial. Weakly expressed subshrub, without sterile shoots. 
Flowering stems, 5—15 in number, ascending, 25—60 cm tall, more or 
less curved, weakly scabrous-setulose in lower part, glabrous and 
smooth above, branched from middle or above. Basal leaves oblong- 
lanceolate, 3-5 mm long, 0.75-1.5 mm broad, distinctly narrowed 
toward base and apex, subobtuse, scabrous-setulose; middle cauline 
leaves narrowly linear-subulate, 7-46 mm long, 0.6—1.5 mm broad, 
somewhat narrowed toward base, cuspidate, with more or less revolute 
margin and veins prominent beneath, all glabrous and smooth; upper 
leaves (in inflorescence) linear-lanceolate, 1.5-4 mm long, 0.25-—0.5 
mm broad, flattened, glabrous and smooth. Inflorescence paniculate, 
fairly spreading. Flowers on very short, 0.25—0.5 mm long, less distinct 
pedicels, 3 together at ends of shortened (0.5—2 mm long) last branches 
of inflorescence. Corolla tubular-infundibuliform, (3.5)4—4.5 mm long; 
lobes more or less reflexed, oblong, obtuse, (1)1.5—1.75 mm long, 
about 0.75 mm broad, usually almost 2 times as short as tube, whitish; 
corolla tube 2.5—-2.75 mm long, scabrous-setulose, like lobes. Anthers 
about 1 mm long. Styles 1.25-1.75 mm long, free at tip. Fruit 2.25— 
2.5 mm finely tuberculate. June to August. 

On calcareous-stony or sandy steppe slopes.— European USSR: 
Black Sea Region (soutwestern part). Endemic. Described from vicinity 
of Nikolaev. Type in Leningrad. 

Note. A quite distinctive species, occupying an intermediate 
position between the heterogeneous species groups Parviflorae and 
Longiflorae; evidently, Kreczetowicz was not aware of its existence. 


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224 


Popov and Chrshanovsky referred it, together with A. bidendata Klok., 
to the above-mentioned hybrid conglomerate. 


25. A. attenuata Klok. sp. n. in Addenda XXII, 698. 

Perennial. Very weakly expressed subshrub without sterile shoots. 
Flowering stems 2—5 in number, suberect or ascending, 26—45 cm tall, 
scabrous-setulose in lower part, weakly scabrous above, branched 
almost from middle or below. Basal leaves oblong-lanceolate or (rarely) 
oblong-elliptic, 2.5—6 mm long, 0.9—1.2 mm broad, broadened at base, 
rarely distinctly narrowed, mucronate, scabrous-setulose along margin; 
middle cauline leaves narrowly linear-subulate to almost setaceous, 
6-32 mm long, 0.5-1.2 mm broad, cuspidate, with revolute margin 
and central vein prominent beneath, more or less scabrous along 
margin; upper leaves (in inflorescence) narrowly linear to linear or 
oblong-lanceolate, 2-7 mm long, 0.3—0.5 mm broad, flattened weakly 
scabrous along margin like floral leaves, latter about 1.5 mm long and 
0.5 mm broad. Inflorescence paniculate, more or less spreading. Flowers 
on pedicels 0.3—1 mm long, 3 together at ends of short branches of 
inflorescence. Corolla tubular-infundibuliform, 3.5—4 mm long; lobes 
more or less recurved, oblong-lanceolate, mucronate, 1.2—1.5 mm long, 
0.5—0.6 mm broad, whitish, scabrous-setulose on outside throughout, 
tube cylindrically narrowed at base, thin. Lobes almost 2 times as 
short as tube, with less distinct or short thickened tip. Anthers about 
1 mm long. Styles 1.2—-1.5 mm long, often unequal, free above middle. 
Fruit about 1.5—2 mm long (immature), fine tuberculate. June to August. 

On calcareous stony slopes.— European USSR: Crimea (vicinity 
of Simferopol). Endemic. Described from Simferopol District. Type 
in Leningrad. 

Note. This species is fairly common in the near and distant (up to 
the village of Partizanskoe) environs of Simferopol, where it is found 
together with A. praevestita Klok., though over lower and more grassy 
sites on the slopes. 


26. A. rumelica Boiss. Diagn. ser. II, 2 (1856) 113.—A. graveolens 
Boiss. Fl. or. III (1873) 41, p.p.; Velen. Fl. Bulg. 238, Suppl. I, 143; 
Grecescu. Consp. fl. Rom. 266; Prodan, Fl. pl. cresc. Rom. I, 968.— 
A. cynanchica var. graveolens Stoian. et Stefan. Fl. Boulg. (1925) 
1044.—A. cynanchica auct. fl. ucr. p.p. 

Perennial. Weakly expressed subshrub. Flowering stems 2-10 in 
number, ascending, 25—70 cm tall, slender, scabrous in lower part and 
often up to middle, upper part glabrous and smooth, highly branched, 
with extremely slender branches. Basal leaves oblong-lanceolate, up 
to 4-5 mm long, and 0.75—1.25 mm broad, scabrous-setulose. Middle 


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225 


cauline leaves narrowly linear-filiform or subulate, 9-38 mm long, 
0.25-0.75 mm broad, cuspidate with more or less revolute margin, 
weakly scabrous; upper leaves (in inflorescence) linear-lanceolate, up 
to 9 mm. long, and about 0.5 mm broad, cuspidate, flattened; floral 
leaves of similar shape, 1-3 mm long, 0.2-0.5 mm broad, smooth. 
Inflorescence paniculate, usually lax. Flowers on very short or more 
distinct, up to 2-3 mm long pedicels. Corolla tubular-infundibuliform 
or sometimes cyathiform, 2—3.5 mm long; lobes weakly deflexed, 
oblong or oblong-ovate, obtuse, 0.8—1.4 mm long, 0.5-0.75 mm broad, 
always considerably, up to one and a half times, shorter than tube, 
whitish, scabrous-setulose; tube, in extreme case, cylindrically narrowed 
at base. Styles 0.4-1.5 mm.long, free almost from base or above 
middle. Anthers up to 1 mm long. Fruit about 1.5 mm long, fine 
tuberculate. June to August. 

On steppe and stony slopes.—European USSR: Bessarabia 
(southern part), Black Sea Region, Lower Don (western part). General 
distribution: Balkans-Asia Minor (Dobrudzha, northern Bulgaria). 
Described from Rumelia. Type in Geneva. 


27. A. graniticola Klok. sp. n. in Addenda XXII, 699.—A. 
tephrocarpa M. Pop. et Chrshan. |.c. quoad plantam maeoticam 
graniticolam, non Czern. sensu proprio.—A. petraea auct. fl. ucr. 
nonnull. non V. Krecz. 

Perennial. Subshrub with woody, thicker (about 2 mm in diameter) 
primary branching axis, forming a mat. Flowering stems ascending, 
10—25(30) cm tall, scabrous-setulose in lower part, middle part 
scabrous, glabrous and smooth above, branched from middle above or 
sometimes below with slender branches. Basal leaves from ovate- 
lanceolate to oblong, 1-4 mm long, 0.5—-0.75 mm broad, obtuse or 
subobtuse, scabrous-setulose, often crowded at base of stem and 
alternately overlapping one another; middle cauline leaves of somewhat 
different shape, linear or linear-spatulate, narrowly linear-subulate and 
linear-lanceolate (upper ones), 8-18 mm long, 0.5—0.75 mm broad, 
rarely broadest above middle, cuspidate, with more or less revolute 
margin and with a vein prominent beneath, glabrous and smooth; upper 
leaves (in inflorescence) oblong-ovate-lanceolate, 2.3 mm long, 0.4— 
0.5 mm broad, acuminate, flat, smooth; floral leaves linear-lanceolate, 
1.25-1.75 mm long, 0.3-0.4 mm broad, narrowly acuminate, smooth. 
Inflorescence more condensed than in other species of this series. 
Flowers 3-5 together subsessile or on extremely short pedicels, at 
ends of branches of inflorescence. Corolla cyathiform-infundibuliform, 
(2.5)2.75—3.25 mm long; lobes more or less deflexed (but not recurved), 
oblong-ovate, obtuse, 1.2—1.3 mm long, 0.6—0.75 mm broad, distinctly 


226 


Ly 
. SWZ 
iN 4 ie D 
it; Y bit Y 
oh #5 ] SF 


| 


iy 
Y D4 
=o 
3 
Plate XIII. 
237 1. Asperula laevissima Klok., upper part of plant, part of inflorescence, flower.— 
2. A. supina M.B., general appearance of plant, part of inflorescence, flower.— 


3. A. savranica Klok., general appearance of plant, part of inflorescence, flower. 


227 


shorter than tube, whitish, scabrous-setulose, corolla tube cuneate, but 
cylindrically narrowed at base. Styles 0.5—1.25 mm long, free above 
the middle. Anthers about | mm long. Fruit about 1.75 mm long, 
finely tuberculate. June to September. 

On outcrops of rubbly rocks (granite, gneiss, hard limestone and 
sandstone).— European USSR: Black Sea Region (southeastern part), 
Lower Don (on border with Black Sea Region). Endemic. Described 
from Azov Sea area (from granite outcrops of ‘Kamennye Mogily’ 
reserve). Type in Kiev. 

Note. Belongs to a petrophilic, ecological group of species similar 
in habit, but having a different origin. Its superficial resemblance to 

240 A. tephrocarpa Czern. or A. petraea V. Krecz., as the resemblance of 
the named species among themselves, is confined to external features. 


Series 6. Vestitae Klok. (s.1.)—Small subshrubs with more or less 
developed sterile shoots, forming mats. Flowering stems 5—35 cm tall, 
branched, usually only in upper part, with few branches. Middle cauline 
leaves narrowly linear to almost setaceous, not longer than 20 mm, 
mostly with revolute margin, floral leaves from oblong- to ovate- 
lanceolate, small or larger, exceeding pistil, always smooth. Flowers 
sessile or on extremely short, less distinct pedicels, 2-3 together at 
ends of branches of inflorescence in fairly dense clusters. Corolla 
very small or larger (2—4.25 mm long); lobes ovate, obtuse or oblong- 
ovate-lanceolate, subobtuse, one and a half times as short as tube, pale 
pink, setulose, rarely pubescent or covered by cuspidate papillae, 
occasionally almost smooth. Style short (0.4-1.5 mm long). Fruit not 
longer than 2.5 mm, finely tuberculate. 

To this series we refer low-montane or premontane Crimean- 
Novorossiisk, xeromorphic species. In the Kreczetowicz system they 
belong to the series Longibracteatae V. Krecz. and Brevibracteatae 
V. Krecz. of the cycle Parviflorae V. Krecz., immediately following 
the series Capitatae V. Krecz., mentioned above. Series Vestitae Klok. 
can be divided, of course, into narrower series, but dividing it into two 
equivalent series is hardly possible. 


28. A. markothensis Klok. sp. n. in Addenda XXII, 700.—A. 
cynanchica §. hirsuta Lipsky in schedis. 

Perennial. Greenish gray subshrub, forming mats. Flowering stems 
suberect, 10-15 cm tall, with up to 3 cm long internodes, densely 
covered with about 0.2 mm long hairs throughout, ternately branched 
above and with axillary flowering branches below. Cauline leaves 
narrowly linear-subulate to almost setaceous, 3.5—-19 mm long, 0.5— 
0.75 mm broad, tapering at top, gently cuspidate, with curved tip, 


241 


228 


margin strongly revolute, densely pubescent with patent hairs up to 0.3 
mm long along margin and on both surfaces; floral leaves oblong- 
lanceolate, 2-4 mm long, 0.5—-1 mm broad, cuspidate or attenuated 
into very short cusp, pubescent. Inflorescence condensed. Flowers 
sessile, 3-7 together at ends of branches. Corolla infundibuliform, 
3.5-4.25 mm long; lobes reflexed oblong-ovate (to almost ovate- 
lanceolate), 1.5-1.75 mm long, about 0.75 mm broad, pink; tube 
cylindrically narrowed at base, pubescent; lobes without thickened 
tip, more or less densely fimbriate along margin, one and a half times 
as short as tube. Styles 0.75-1.25 mm long, free at top. Anthers about 
1 mm long. Ovary weakly tuberculate. Fruit not known. June to August. 

Mountain slopes.— Caucasus: Western Transcaucasia (Markotkh 
Range near Novorossiisk). Endemic. Described from the indicated 
locality. Type in Leningrad. 


29. A. lipskyana V. Krecz. in Grossh. Fl. Kavk. IV (1934) 26; 
Grossh. Opred. rast. Kavk. 248. 

Perennial. Glaucescent-green subshrub, forming lax mats. 
Flowering stems 10-35 cm tall, with up to 4 cm long internodes, 
diffusely scabrous-setulose in lower part, glabrous and smooth above, 
furcate or ternately branched at top, not densely foliated. Basal leaves 
oblong, about 2 mm long and 0.5 mm broad, obtuse. Cauline leaves 
narrowly linear or (upper ones) narrowly linear-lanceolate, up to 17- 
21 mm long, 0.5-1.25 mm broad, cuspidate, flat or with slightly 
revolute margin, weakly scabrous-setulose along margin, mainly at 
base; floral leaves linear or oblong-lanceolate, very narrow and almost 
hairy at apex, 2-5 mm long, 0.2-1 mm broad. Inflorescence fairly 
spreading with divaricate branches. Flowers 3 or more together on 
ultimate branches on inflorescence almost sessile or with extremely 
short, less distinct pedicels. Corolla infundibuliform, 2.5—3.5 mm long; 
lobes strongly reflexed, oblong-ovate up to 1.5 mm long and 0.75 mm 
broad, pink; tube with cylindrical base, weakly scabrous-setulose; lobes 
glabrous and smooth, with less distinct whitish thickened tip. Style up 
to 1.5 mm long. Fruit 2—2.5 mm long, finely tuberculate. June to 
August. 

On limestone outcrops in lower mountain belt.— Caucasus: 
Ciscaucasia (in northwestern part, along Urup River), western 
Transcaucasia (vicinity of Novorossiisk and Gelendzhik). Endemic. 
Described from vicinity of Novorossiisk (crags near cement factory). 
Type in Leningrad. 

Note. We do not consider it possible to unite this species with the 
preceding one, expanding its limits indefinitely, as proposed by A. 
Kolakovsky in his notes to the exsiccatae of ““Gerbariya Flory SSSR” 


242 


229 


under the number 3696 (specimens under this number represent a 
mixture of these species). Both species differ not only in pubescence, 
but their geographical distribution is undoubtedly different. The 
occurrence of intermediate forms in areas of contact (by the way, we 
did not find such forms in herbaria) cannot eliminate, of course, the 
evident differences between the two geographic races. 


30. A. aemulans V. Krecz. in manuscr.; Klok. in Addenda XXII, 
700.—A. supina auct. fl. taur. pro min. parte, non M.B. 

Perennial. Bluish green subshrub with numerous, densely foliated 
sterile shoots, forming dense mats. Flowering stems suberect or 
ascending, 7-20 cm tall, with up to 1.5—4.5 cm long internodes, fairly 
slender, lower part weakly scabrous-setulose and densely foliated with 
shortened axillary branches, upper part glabrous and smooth, weakly 
furcate or ternately branched. Basal leaves oblong-ovate, to oblong- 
lanceolate, 1-2 mm long, 0.5-0.75 mm broad, obtuse or somewhat 
mucronate, glabrous and smooth along margin, middle cauline leaves 
narrowly linear-subulate, 5—20 cm long, 0.25—0.75(1) mm broad, 
cuspidate or often attenuated into extremely short soft mucro, scabrous- 
setulose along margin with broad vein which is not prominent beneath 
and extremely narrow bands of tissue on its sides, almost appressed to 
stem or slightly patent; upper leaves (below inflorescence and at base 
of its main branches) linear-lanceolate, about 1 mm broad, shorter 
than middle leaves; floral leaves oblong or ovate-lanceolate,.2-4 mm 
long, 0.5-1.25 mm broad, narrowly acuminate or attenuated into a 
cusp, margin smooth. Inflorescence more or less condensed. Flowers 
sessile, (3)5—9, gathered at ends of branches and forming dense rarely 
almost capitate clusters. Corolla infundibuliform, 3.5—4 mm long; lobes 
recurved, oblong-ovate, obtuse or oblong-ovate-lanceolate, subobtuse, 
1.5-1.75 mm long, 1—1.25 mm broad, pale pink, without distinct whitish 
thickened tip; corolla tube entirely glabrous and smooth or sparsely 
setulose at base; lobes smooth. Styles 0.4-0.8 mm long, free almost 
up to base. Anthers 1-1.25 mm long. Ovary finely tuberculate. Fruit 
unknown. June to August. 

Mountain slopes.—European USSR: Crimea (southern slopes of 
mountains in the regions of Alushta, Sudak, Kokkoz). Endemic. 
Described from vicinity of Alushta (southwestern slope of Chatyr- 
Dag in a preserve). Type in Leningrad. 

Note. Kreczetowicz wavered in acknowledging the separateness 
of this species on account of its similarity to A. caespitans Juz. (A. 
supina V. Krecz. pro max. parte, non M.B.). However, it cannot be 
placed in a single series with the last-named species because of the 


243 


230 


extremely profound, though not easily visible, differences between 
them. 


31. A. praevestita Klok. sp. n. in Addenda XXII, 701. 

Perennial. Grayish green subshrub, with more or less developed, 
densely foliated sterile shoots. Flowering stems suberect or ascending, 
7-35 cm tall, with up to 2-3.5 cm long internodes, slender, in lower 
half densely pubescent with distinct patent and retrorse, setaceous, 
hairs about 0.2—0.3 mm or sometimes even 0.5 mm long (f. villosissima 
Klok.), in inflorescence glabrous and smooth or only in its lower 
internodes branched, very mildly scabrous-setulose, above the middle 
furcately and ternately branched. Basal leaves from oblong-ovate to 
oblong-lanceolate, 1-4 mm long, 0.5—1 mm broad, somewhat obtuse 
or mucronate, glabrous and smooth or slightly scabrous along margin; 
cauline leaves narrowly linear or uppermost linear-lanceolate, to almost 
subulate or setaceous, 3-16 mm long, 0.3-1 mm broad, slightly 
narrowed at apex, cuspidate, with thickened and more or less revolute 
margins and thick, whitish vein prominent beneath, basal and cauline 
leaves pubescent, with hairs up to 0.2 mm long, upper leaves setulose 
along margin, uppermost leaves glabrous and smooth; floral leaves 
oblong or almost ovate-lanceolate, 1-2.5 mm long, 0.25—1 mm broad, 
cuspidate, glabrous, smooth. Inflorescence more or less condensed, 
usually with 4-7 main branches. Flowers on its terminal branches 
grouped in clusters of 3-7, sessile or almost sessile. Corolla 
infundibuliform, 2,5-3.5 mm, often about 3.5 mm long; lobes more or 
less deflexed ovate, nearly 1.5 mm long, 0.75—1 mm broad; corolla 
tube comparatively broader, densely setulose; lobes diffusely setulose 
along margin, with very small, whitish, thickened tip. Style 0.25—0.75 
mm long. Fruit about 1.75 mm long covered with cuneate tubercles. 
June to August. 

On = limestone-marly outcrops.—European USSR: Crimea 
(Tarkhankut Peninsula, vicinity of Simferopol, Sevastopol and 
Eupatoria). Endemic. Described from Eupatoria District (Dzhan-Gul 
area). Type in Leningrad. 

Note. According to the observations of N.M. Chernova on the 
Tarkhankut Peninsula, the corolla of this species is light blue. The 
color is completely lost, however, in dried materials. In the vicinity of 
Simferopol, the flowers are white. The Tarkhankut plant is referred to 
as f. villosissima Klok., and probably represents a distinct species, 
requiring additional study. 


32. A. vestita V. Krecz. in Grossh. Fl. Kavk. IV (1934) 15, quoad 
nominem. excl. plantam caucasicam.—A. littoralis D’urv. Enum. 15, 


244 


231 


sec. Ldb. Fl. Ross. II, 1 (1844) 399, non Sibth.—Exs.: Herb. norm ed. 
ab. J. Dérfler, No. 3944 (sub A. cretacea Willd.). 

Perennial. Glaucescent-pale green subshrub, with more or less 
developed sterile shoots, forming lax mats. Flowering stems, ascending, 
10-30 cm tall, with up to 3.5 cm long internodes, arcuate at base, 
straight above, slender (0.5—0.75 mm thick), weakly scabrous-setulose 
in lower part; glabrous and smooth above, furcately or often ternately 
branched in inflorescence. Basal leaves linear-oblong, 2-4 mm long, 
0.5—0.75 mm broad, approximately of same length as corresponding 
shortened internode or considerably longer, obtuse on mucronate. 
Cauline leaves (up to inflorescence) linearly-setaceous, 5-18 mm long, 
more or less about 0.5 mm broad, tapered at top, cuspidate, with 
nonrevolute margin; lower leaves slightly longer than corresponding 
internodes, the remaining leaves 2—3 times as short; leaves on sterile 
shoots several times as long as internodes, lower and middle cauline 
leaves, like leaves on sterile more or less distinctly scabrous setulose 
along margin, upper leaves (in inflorescence) linear-lanceolate, up to 
1 mm broad, attenuate-cuspidate, smooth along margin; floral leaves 
oblong- or almost ovate-lanceolate, very small, 1-1.5 mm long, 0.2— 
0.5 broad. Inflorescence with few branches, separating at acute angle. 
Flowers grouped in clusters of 3—7 on terminal branches, sessile or on 
less distinct extremely short, not longer than 0 .5 mm pedicels. Corolla 
infundibuliform, 2.5—-3 mm long; lobes more or less recurved, ovate 
or oblong-ovate, 1—1.25 mm long, 0.5-0.75 mm broad pale pink; tube 
comparatively broad, densely setulose; lobes scabrous-setulose along 
margin, with less distinct thickened tip. Style 0.5-1 mm long. Fruit 
about 2 mm long, densely covered with obtusely cuneate tubercles. 
June to August. 

On rubbly limestone mountain slopes and sandy-shell-rock banks 
of brackish lakes.— European USSR: Crimea (Karadag, vicinity of 
Feodosia and Sudak). Endemic. Described from vicinity of Karadag. 
Type in Leningrad. 

Note. The type of species was selected by Kreczetowicz from 
Crimean material (annotated in the herbarium and cited in manuscript) 
was not cited in the original description in ‘Flora Kavkaza.” In 
Caucasian material, no specimens of this species were found; in all 
probability it does not occur there. Identification of this species with 
A. cretacea Willd. (apud Roem. et Schult. Syst. Veg. III (1818) 329) 


‘remains somewhat of puzzle. Mention of A. cretacea Willd. as a species 


is absent.in the manuscript of Kreczetowicz; it is discussed only in 
connection with the foregoing, cited exsiccatae. At present we are 
unable to confirm the Crimean species to which this name can be 


245 


232 


applied. The epithet “cretacea” evidently fits only A. supina M.B. 
(1808). 


33. A. infracta Klok. sp. n. in Addenda XXII, 701. 

Perennial. Yellowish or grayish, green subshrub. Flowering stems 
at base almost decumbent, above bent at an angle and ascending in an 
arched or zig-zag manner, 15-25 mm tall, with up to 2-3 mm long 
internodes, comparatively thick (1-1.5 mm thick at bottom), with fairly 
thick, setulose pubescence in lower part, glabrous and smooth above, 
in inflorescence ternately or in part furcately branched. Basal leaves 
from ovate-deltoid, obtuse, to oblong-lanceolate, more or less 
acuminate, 1-4 mm long, about 1 mm broad, longer than corresponding 
shortened internodes, glabrous and smooth. Cauline leaves from 
narrowly linear to linear-lanceolate (upper ones), 3-14 mm long, 0.5— 
1.5 mm broad (at base), cuspidate or attenuated into a short mucro, 
with more or less revolute margin, somewhat fleshy, bent almost at 
right angles from the stem and curved in an arch at bottom; basal and 
middle leaves scabrous-setulose along margin, upper ones smooth; floral 
leaves oblong or ovate-lanceolate, 1.5—2.5 mm long, 0.3—1 mm broad, 
cuspidate, smooth. Inflorescence lax, with comparatively numerous 
patent branches, flowers grouped in clusters of 3-7 on terminal 
branches, sessile. Corolla infundibuliform, 2—2.5 mm long; lobes ovate, 
recurved, 0.75—1 mm long and up to 0.75 mm broad, pale pink; tube 
comparatively broad, not densely setulose; lobes diffusely setulose- 
serrulate along margin, with whitish thickened tip. Styles 0.5-1 mm 
long, free above the middle. Anthers about 1 mm long. Ovary finely 
tuberculate. Fruit unknown. June to August. 

On sandy-shell-rock deposits and in absinthe-herbaceous steppe.— 
European USSR: Crimea (Tarkhankut Peninsula, vicinity of Eupatoria). 
Endemic. Described from vicinity of Eupatoria. Type in Leningrad. 


34. A. praepilosa V. Krecz. in manuscr.; Klok. in Addenda XXII, 
702.—A. cynanchica var. affinis Wulf in Zap. Krym. obshch. estestv. 
XI (1929) 77, ex p. non A. affinis Boiss. and Huet. 

Perennial. Slightly pale green subshrub, with more or less 
developed densely foliated sterile shoots, forming fairly lax mats. 
Flowering stems suberect, 10-20 mm long with internodes usually not 
longer than 1.5 cm, slender (0.5-0.75 mm thick), fairly densely 
pubescent with short, up to 0.2-0.25 mm long, thin hairs throughout, 
comparatively densely foliated, with shortened sterile shoots in axils 
of leaves, furcate or ternately branched in upper part. Basal leaves 
from oblong-ovate-lanceolate subobtuse to linear-oblong, cuspidate, 
1-3 long, 0.1-0.75 mm broad, fairly weakly scabrous-setulose along 


246 


233 


margin, considerably longer than corresponding highly shortened 
internodes; cauline leaves narrowly linear, 5-15 mm long, 0.5—1 mm 
broad, cuspidate or attenuated into distinct apical mucro, margin 
revolute, along margin and whole surface pubescent, with up to 0.4 
mm long hairs, vein fairly thin and weakly prominent beneath, somewhat 
patent and curved, longer and only at top somewhat shorter than 
corresponding internode; floral leaves oblong or almost ovate- 
lanceolate, 1-3 mm long, 0.25-0.75 mm broad, cuspidate, glabrous, 
slightly scabrous along margin, rarely scabrous-ciliate. Inflorescence 
condensed. Flowers 3—5 together at end branches in small dense 
clusters, sessile or subsessile. Corolla infundibuliform, 2.5—3 mm long, 
with deflexed oblong-ovate about 1.25 mm long and 0.5 mm broad, 
pale pink, with whitish thickened tip, diffusely papillose along margin. 
Style about 0.3-0.5 mm long, free up to almost base. Ovary finely 
tuberculate. May to June. 

On rocky mountain slopes.— European USSR: Crimea (Kerch 
Peninsula, Mt. Opuk). Endemic. Described from the indicated locality. 
Type in Leningrad. 

Note. This species, along with the following one, belongs to a 
more narrow group, Cimmericae Klok., a component of an extensive 
series Vestitae Klok. 


35. A. cimmerica V. Krecz. in manuscr.; Klok. in Addenda XXII, 
702.—A. cynanchica var. affinis Wulf in Zap. Krymsk. obshch. est. 
XI (1929) 77 ex p. non A. affinis Boiss. and Huet. 

Perennial. Glaucescent-green or dark green subshrub with 
numerous, densely foliated, sterile shoots, forming small, but dense, 
pulvinate mats. Flowering stems angularly suberect or obliquely 
ascending, 4-14 cm tall, with up to 1-1.5 cm long internodes, fairly 
slender (0.5-—0.75 mm thick), scabrous-setulose throughout, more 
weakly so above, comparatively densely foliated, with shortened sterile 
twigs in axils of lower and middle leaves, furcate or ternately branched 
in upper part. Basal leaves oblong-lanceolate, about 1-2 mm long, 
0.75 mm broad, obtuse, glabrous and smooth along margin; cauline 
leaves narrowly linear, 3-10 mm long, 0.3—1 mm broad, at top slightly 
contracted (more distinctly in upper leaves), cuspidate, often with 
somewhat revolute margin, scabrous-setulose along margin, with broad 
vein, mildly prominent beneath, and extremely narrow bands of tissue 
along its sides, more or less patent and arcuate; floral leaves oblong- 
lanceolate, 1.5-3 mm long, 0.5—0.75 mm broad, narrowly acuminate, 
scabrous-setulose along margin also. Inflorescence with shortened 
branches. Flowers 3-7 in dense clusters, sessile or almost sessile, 
aggregated at ends of secondary branches. Corolla infundibuliform, 


247 


234 


3-3.5 mm long; lobes deflexed, ovate or oblong-ovate, 1.25-1.5 mm 
long, 0. 75-1 mm broad, pale pink, with very small, whitish, callous 
tips; corolla tube slightly fairly thinly narrowed at base, densely 
scabrous-setulose throughout; lobes setulose along margin also. Styles 
0.5-1 mm long, free almost up to base. Fruits about 1.5 mm long, 
covered with hemispheroidal tubercles. May to July. 

On limestone rocks and littoral shell-rock sands.— European USSR: 
Crimea (Kerch Peninsula). Endemic. Described from dunes along Sea 
of Azov on Kerch Peninsula. Type in Leningrad. 


Series 7. Pedicellatae Klok.—Differs from the preceding series in 
lax inflorescence and flowers on conspicuous pedicels. Characteristic 
feature comprises subglabrous corolla, which is also observed in 
Vestitae Klok. (in A. aemulans V. Krecz.). 

To this series we refer two Ciscaucasian mountane (but not alpine) 
species. Of these, A. biebersteinii V. Krecz. was referred by the author 
[Kreczetovicz] to the series Leianthae V. Krecz. (see characteristics of 
series Petreae Klok.). 


36. A. biebersteinii V. Krecz. in Grossh. Fl. Kavk. IV (1934) 26; 
Kolak. Fl. Abkh. IV, 130.—A. supina M.B. FI. taur.-cauc. I (1808) 
101 ex p. quoad pl. caucasicam.—A. cynanchica auct. Fl. cauc. p.p. 
non L.—A. vestita V. Krecz. in Grossh. Fl. Kavk. IV (1934) 125, p.p. 
quoad pl. caucasicam. 

Perennial. Pale green subshrub with weakly developed, fruitless 
shoots. Flowering stems ascending or suberect, to almost erect, 3.5—30 
cm tall, with up to 1-4.5 cm long internodes, fairly weakly scabrous- 
setulose in lower part, glabrous and smooth above, furcately and 
ternately branched above the middle; sparsely or very densely foliated. 
Basal leaves oblong-ovate (obovate), very small, 1—-1.5 mm long, 0.5— 
0.7 mm broad, cuspidate; middle cauline leaves narrowly linear to 
almost setaceous, 3—25(30) mm long, 0.3—1(1.5) mm broad, cuspidate, 
with strongly revolute margin, more or less scabrous-setulose along 
margin; floral leaves linear or oblong-lanceolate, 14.5 mm long, 0.2— 
0.75 mm broad. Inflorescence lax. Flowers aggregated in threes at 
ends of last branches; middle flowers of the cyme on less noticeable 
pedicel, lateral flowers on 0.5-—2 mm long pedicels. Corolla 
infundibuliform, 2.75—3.75 mm long; lobes deflexed oblong-ovate, 1- 
1.75 mm long, 0.5—0.8 mm broad, pale pink, with conspicuous, whitish, 
callus tip; corolla tube comparatively broad, glabrous, with isolated 
tubercles in lower part or, rarely, slightly setulose. Anthers about 1 
mm long. Styles 0.5—1 mm long, free at tip. Ovary finely tuberculate. 
Fruits 1.5-2 mm long, almost globose or short ellipsoidal, finely 
tuberculate. June to August. 


235 


Mountain slopes up to subalpine belt.— Caucasus: Ciscaucasia 
(western part). Described from vicinity of Pyatigorsk, Mt. Goryachaya. 
Type in Leningrad. 


37. A. pedicellata Klok. sp. n. in Addenda XXII, 703. 

Perennial. Grayish green subshrubs with numerous, densély foliated 
sterile shoots, forming mat. Flowering stems suberect, 5-8 cm tall, 
with up to 1 cm long internodes, scabrous (from dense or in middle 
part of stem scattered, acutely cuneate, very fine setae) throughout, 
ternately or furcately branched in upper part and with more or less 
developed, short sterile axillary twigs densely foliated. Leaves narrowly 
linear or linear-spatulate, broadest in the upper quarter of stem (middle 
cauline), 2.5-10 mm long, 0.25—0.75(1) mm broad, acuminate or 
apiculate, but without attenuated cusp at apex, flat, with somewhat 
revolute margin, margin scabrous-setulose (setae inclined forward, 
toward apex of leaf), leaves on stems and sterile shoots more or less 
spreading and curved downward in a crescent; floral leaves linear, 
acuminate or linear-lanceolate, 3-4.5 cm long, 0.5—0.75 mm broad. 
Inflorescence fairly lax. Flowers on 0.5—2 mm long scabrous pedicels, 
aggregated in threes on terminal branches. Corolla _ tubular- 
infundibuliform, 4.5—5 mm long; lobes reflexed, oblong, ligulate, 1.75— 
2 mm long, 0.7—0.8 mm broad, pale pink, with whitish, callous tip; 
corolla tube covered with scattered, very fine, cuneate setae, visible 
under a magnifying glass. Styles 1.15-1.75 mm long, free only at tip. 
Ovary and fruit densely covered with very fine obtuse tubercles. June 
to August. 

On limestone rocks.— Caucasus: Ciscaucasia (Mt. Beshtau), 
Dagestan. Endemic. Described from the summit of Mt. Beshtau near 
Pyatigorsk. Type in Leningrad. 

Note. Akinfev’s specimens from Mt. Beshtau were referred by 
Kreczetowicz to A. biebersteinii V. Krecz.; however, the plant turned 
out to be quite different from the aforementioned species in most 
characteristics. It was closer to A. cristata (Somm. and Lev.) V. Krecz., 
in general appearance but differed sharply from the latter species also 
in most respects. 


Series 8. Cristatae Klok.— Small subshrubs with developed sterile 
shoots, forming mat. Flowering stems 3-20 cm tall. Middle cauline 
leaves mostly narrowly linear, flat or with slightly revolute margin. 
Floral leaves linear or oblong-lanceolate, rarely ovate-lanceolate, up 
to 4—5 mm long and up to 0.75-1.25 mm broad, exceeding fruit, 
entire. Inflorescence compact, almost corymbose. Flowers sessile or 
subsessile, 3-7 together on terminal branches in fairly dense or more 


249 


236 


lax clusters. Corolla tubular-infundibuliform, comparatively large, 4.5— 
8 mm long; lobes oblong-ovate, obtuse lobes, one and a half to two 
times as short as tube, glabrous or subglabrous, rarely puberulent, 
pink, with whitish, callous tips. Anthers oblong, 0.75-1.25 mm long. 
Style 0.5—3 mm long. Fruits 1.5-2.5 mm long, covered with fine, 
obtuse tubercles. 

In this series we include the alpine species of Ciscaucasia and 
Dagestan. The species of this series known to Kreczetowicz were 
referred by him to the cycle Pontigenae V. Krcz. and series 
Brachyphyllae V. Krecz., which, as a result, completely corresponds 
to our series, differing only in name. We have changed the name of the 
series, following the accepted convention of basing the names of the 
series on the type species. 


38. A. dasyantha Klok. sp. n in Addenda XXII, 704. 

Perennial. Grayish dark green subshrub, with elongated slender 
sterile shoots, forming lax mats. Flowering stems ascending or semi- 
decumbent, 10-15 cm tall (long), with up to 2.5-3.5 cm long 
internodes, fairly densely puberulent with separate thin very short 
hairs throughout, ternately or furcately branched in upper part, with 
more or less developed sterile axillary branches below, not densely 
foliated. Lower leaves oblong-lanceolate, about 1 mm long and 0.5 
mm broad, cuspidate, middle cauline leaves narrowly linear, acuminate 
to narrowly linear-lanceolate; 7-15 mm long, 0.8-1 mm broad, 
cuspidate and usually tapered at apex into very short, soft cusp; upper 
leaves (at base of inflorescence) linear-oblong to linear-lanceolate, 5- 
13 mm long, 1—1.5 mm broad, also usually with cusp; floral leaves 
from narrowly linear to ovate-lanceolate, 2.5 mm broad; leaves on 
sterile shoots shorter and distinctly narrower, 2.5-10 mm long, 0.25-— 
0.75 mm broad; all leaves flat with scarcely revolute margin, more or 
less patent and curved, pubescent with fine hairs along margin and 
midrib above and beneath. Inflorescence corymbose, fairly dense. 
Flowers sessile or subsessile, aggregated in threes usually on very 
short but varied, terminal, branched inflorescences. Corolla tubular- 
infundibuliform, 5.5—7 mm long; lobes more or less deflexed, oblong- 
ovate, 1.5-2.5 mm long, 0.75-1.226 mm broad, pale pink, with less 
noticeable whitish, callous tip; corolla tube thicker, puberulent; lobes 
distinctly barbate along margin at tip. Anthers 0.75—1 mm long. Style 
2—2.5 mm long, free at tip. Ovary finely tuberculate. Fruits unknown. 
June to August. 

On rocks in alpine zone.— Caucasus: Ciscaucasia (Ingush Alps). 
Endemic. Described from Ingush Alps (Dzharesht, southeast from 
Ordzhonikidsze). 


250 


237 


39. A. accrescens Klok. sp. n. in Addenda XXII, 704. 

Perennial. Glaucescent-green subshrub with more or less developed 
sterile shoots, forming lax mats. Flowering stems ascending, 10-20 
cm tall, angularly curved in lower part, with up to 3.5-5 cm long 
internodes, not very slender, glabrous, or weakly scabrous (because of 
acute retrorse cuneate papillae nearly unnoticeable in profile) 
throughout, ternately branched above with more or less developed 
sterile axillary twigs; cauline leaves linear or narrowly linear, 3—20 
mm long, 0.5-1.5 mm broad (middle ones not shorter than 1 cm), 
cuspidate and mostly tapered into very short, soft cusp, flat or nearly 
so, scabrous-setulose, rarely weakly scabrous along margin; floral 
leaves mostly linear- or oblong-lanceolate, 2.5-5 mm long, 0.5-1.5 
mm broad. Flowers in clusters of 3-7, aggregated on ends of branches 
of almost corymbose inflorescence, sessile or on very short, less distinct 
pedicels. Corolla tubular-infundibuliform, 5—8 mm long; lobes reflexed 
on ovate or oblong-ovate, 2-2.5 mm long, 1—1.25 mm broad, pink, 
with fairly large, whitish, callous tips; corolla tube fairly slender, 
entirely glabrous and smooth like lobes. Anthers 1-1.25 mm long. 
Styles 1.5-3 mm long, unequal, free only at tip. Ovary finely 
tuberculate. Fruits unknown. June to August. 

On rocks and glacial moraines in alpine and subalpine zone.— 
Caucasus: Ciscaucasia. Endemic. Described from peak of Mt. Atsgara 
in foothills of Peredovoi Range of the Great Caucasus Range. Type in 
Leningrad. 

Note. In working over the materials in Leningrad, Kreczetowicz 
referred specimens of this species partly to A. cristata (Somm. et 
Lev.) V. Krecz., and partly to A. biebersteinii V. Krecz. Our species 
differs sharply from the latter in having considerably broader and 
flatter, not setaceously convoluted cauline leaves, a more compact 
inflorescence with more developed floral leaves and a much larger, 
glabrous corolla. From A. cristata (Somm. et Lev.) V. Krecz., it differs 
by the comparatively large size of the plant and, particularly by the 2— 
3 times longer middle cauline leaves. The stems, even when identical 
in length, are much thicker (+1 mm instead of +0.3 mm) and are 
completely glabrous, whereas in A. cristata (Somm. et Lev.) V. Krecz. 
they are covered with distinct setae, seen only under a magnifying 
glass. It also differs no less sharply from all other known species. It 
is fairly widespread in the mountains of the Greater Caucasus Range, 
as compared to A. cristata (Somm. et Lev.) V. Krecz. 


40. A. cristata (Somm. et Lev.) V. Krecz. in Grossh. Fl. Kavk. IV 
(1934) 27.—A. cynanchica var. cristata Somm. et Lev. in Tr. SPb. 
bot. sada (1900) 204.—A. cynanchica auct. fl. cauc. p.p. non L. 


253 


254 


238 


Perennial. Green subshrub, with numerous sterile shoots, forming 
mat. Flowering stems more base angularly suberect, 5-10 cm tall, 
with up to 1-2 cm long internodes, fairly slender (about 0.5 mm 
thick), scabrous-setulose throughout, ternately branched in upper part, 
lower part often with developed fruitless axillary twigs, comparatively 
densely foliated. Lower leaves from oblong-ovate to linear-oblong, 1- 
2.5 mm long, 0.3-0.8 mm broad, smooth or weakly scabrous along 
margin; cauline leaves from narrowly linear-spatulate (lower ones) to 
linear-lanceolate, mostly narrowly linear, 3-10 mm long, 0.5—1.25 
mm broad, cuspidate, flat, scabrous-setulose along margin, floral leaves, 
0.25-—0.75 mm broad narrowly linear or narrowly linear-lanceolate. 
Inflorescence almost corymbose. Flowers 3, together at ends of few 
branches, rarely 7, in lax clusters, almost sessile or on extremely short 
(about 0.5 mm long) pedicels. Corolla tubular-infundibuliform, 5—7 
mm long; lobes deflexed, oblong-ovate, 1.5—2 mm long, 0.8-1.25 mm 
broad, pink, with whitish callous tips; corolla tube slender, with 
scattered or solitary, fine setae in lower part; lobes smooth along 
margin. Anthers about 1 mm long. Styles 1-2.5 mm long, somewhat 
unequal, free in upper part. Fruits 1.5—2.25 mm long, finely tuberculate. 
June to August. 

On rocks and glacial moraines in alpine zone.— Caucasus: 
Ciscaucasia. Endemic. Described from Mt. Elbrus. Already known 
from the moraines of Mizhirgi Glacier. Type in Florence? 


41. A. alpina M.B. FI. taur.-cauc. II (1819) 103; DC. Prodr. IV, 
584; Kreczetowicz in Grossh. Fl. Kavk. IV, 26.—C. cynanchica B. 
alpina C.A. Mey. Verzeichn. Pfl. Cauc. (1830) 52; Boiss. Fl. or. III, 
40 ex p.; Lipsky in Tr. Tifl. bot. sada, IV, 333, p.p. 

Perennial. Grayish-green subshrub, with numerous sterile shoots, 
forming fairly dense mats. Flowering stems suberect, mostly 10—15 
mm tall, slender (0.3-0.5 mm thick), with up to 1-2.5 mm long 
internodes, throughout scabrous from acute cuneate papillae or 
scabrous-setulose, rarely almost smooth above, ternately branched in 
upper part, lower part with more or less developed, sterile, axillary 
twigs. Basal leaves linear-oblong to narrowly elliptic, 1.5—3 mm long, 
0.5-0.75 mm broad, distinctly narrowed at base, mucronate, margin 
smooth; cauline leaves narrowly linear, S-12 mm long, 0.5—0.75 mm 
broad, with more or less distinctly revolute margin, not densely 
scabrous-setulose to serrulate (in inflorescence) along margin; lower 
leaves distinctly narrowed at base, slightly broadened toward apex, 
mucronate; upper leaves cuspidate, in inflorescence narrowly linear- 
lanceolate, sometimes up to 1 mm broad; floral leaves linear- or oblong- 
lanceolate, 3-4.5 mm long, 0.25-0.75 mm broad, fairly narrowly 


239 


Plate XIV. 


251 1. Asperula paniculata Bge., general appearance, unfolded corolla, fruit.—2. 
A. karataviensis Pavl., upper part of plant, whorl of middle cauline leaves, 
unfolded corolla, pistil, fruit from upper portion.—3. A. octonaria Klok., upper 

part of plant, unfolded corolla, pistil, fruit. 


255 


240 


acuminate, often distinctly narrowed at base, sometimes almost elliptic, 
nearly flat, weakly or more distinctly scabrous-setulose along margin; 
rarely few leaves rudimentary, almost filiform. Inflorescence fairly 
compact, almost corymbose. Flowers 2(3) together on terminal, 
shortened branches; middle flowers of clusterlike cymes almost sessile, 
lateral flowers on very short, but somewhat distinct, 0.5—0.75 mm 
long, pedicels. Corolla tubular-infundibuliform, 4.5—6 mm long; lobes 
spreading with recurved tip and oblong-ovate, 2—2.5 mm long, 0.75- 
1 mm broad, pale pink with less distinct whitish, callous tips; corolla 
tube not very slender, with scattered or isolated acute papillae in lower 
part; lobes smooth along margin. Anthers 1-1.25 mm long. Styles 1- 
2 mm long, free at tips. Fruits 1.5—2.5 mm long, covered with fine 
vesicular tubercles. June to August. 

On mountain slopes and stony screes in alpine and subalpine 
zone.—Caucasus: Dagestan, eastern Transcaucasia. Described from 
Dagestan. Type in Leningrad. 


42. A. intersita Klok. sp. n. in Addenda XXII, 705. 

Perennial. Glaucescent-pale green subshrub, with fairly densely 
foliated sterile shoots, forming mats. Flowering stems obliquely | 
ascending or suberect, 3—12 cm tall, with up to 1-3 cm long internodes, 
slender (not thicker than 1 mm), glabrous and smooth throughout, 
weakly furcately or ternately branches in upper part, not densely 
foliated. Leaves dissimilar in shape and size (heterophylly), 2-12 mm 
long, 0.3—1 mm broad, flat or nearly so, with scarcely revolute margin, 
glabrous and smooth even along margin, and only basal leaves weakly 
scabrous at base; all cauline leaves more or less declinate from stem 
and somewhat curved, lowest leaves on stem narrowly linear-lanceolate, 
acuminate, very small; following these (lower and middle) narrowly 
spatulate, widest in lower quarter and toward base narrowed into 
petiole, mucronate, leaves situated higher (upper-middle) narrowly 
oblong-lanceolate, cuspidate, floral leaves oblong or ovate-lanceolate, 
2.5—4 mm long, 0.5—1.25 mm broad, attenuated cuspidate. Inflorescence 
condensed, weakly branched, consisting of 1-4 clusters with 3-7 
flowers in each cluster, at ends of branches. Flowers almost sessile or 
on very short (0.5 mm long) less distinct pedicels. Corolla 
infundibuliform, 4.5—5 mm long, lobes more or less deflexed, oblong- 
ovate, 1.5-2 mm long, 0.75—-1.25 mm broad, pale pink, with fairly 
large whitish, callous tip, completely glabrous and smooth. Anthers 
oblong, 1-1.25 mm long. Styles free almost up to base, ovary finely 
tuberculate. Fruit about 2 mm long and broad, subglobose, covered 
with fine obtuse tubercles. July to August. 


256 


241 


On mountain slopes in alpine zone.—Caucasus: Ciscaucasia (along 
boundary with western Transcaucasia). Endemic. Described from Mt. 
Fisht in Cherkessia. Type in Leningrad. 

Note. This species differs sharply from other representatives of 
the series, being closer to A. abchasica V. Krecz. (from series Ponticae 
Klok.). In the latter, however, the leaves are longer and considerably 
narrower, with the margins strongly revolute and sparsely but distinctly 
setulose; the corolla also is longer and with a narrower tube; the stem 
is considerably thinner, with sparse, small setae in the lower part. 
Occupying, to some extent, an intermediate position between the two 
series, at the same time A. intersita Klok. differs by its characteristic 
heterophylly from all members of both series. 


Series 9. Ponticae Klok.—Low subshrubs with numerous sterile 
shoots, forming dense mats. Flowering stems 3-10 cm tall. Floral 
leaves oblong or often ovate-lanceolate, up to 4-5 mm long and 1.5 
mm broad. Inflorescence of 1-3 dense almost capitate clusters, with 
3-9 flowers in each cluster. Flowers sessile or on less distinct, very 
short pedicels. Corolla tubular-infundibuliform, 5—5.5 mm long; lobes 
oblong-ovate, one and a half times as short as tube, glabrous and 
smooth or very weakly scabrous-setulose, pink. Style 1.5—2 mm long. 
In other respects, they resemble the preceding series. 

In this series we include the western Transcaucasian species of 
the series Dolichophyllae V. Krecz. of cycle Pontigenae V. Krecz., 
established by Kreczetowicz. 


43. A. abchasica V. Krecz. in Grossh. Fl. Kavk. IV (1934) 127; 
Kolak. Fl. Abkh. IV, 129.—A. cynanchica var. alpina Alb. Prodr. FI. 
Colch. (1895) 121.—Ie.: Kolak. l.c. tabl. X.—Exs.: GRF, No. 3238. 

Perennial. Grayish-green, light colored subshrub with numerous 
sterile shoots, forming fairly dense mats; flowering stems suberect at 
base, 3.5—10 cm tall, with up to 1-2.5 cm long internodes, slender 
(about 0.5 mm thick), scabrous setulose below nodes in lower part 
and above, furcately or ternately branched upwards or sometimes from 
middle, comparatively densely or more sparsely foliated. Cauline leaves 
from linear-oblong (the lowest) to narrowly linear, almost setaceous 
(middle ones, mostly) and very narrowly linear-lanceolate (in 
inflorescence), 2.5-17 mm long, 0.5-1 mm broad, cuspidate, with 
more or less revolute margin, lower part of margin scabrous, or very 
weakly scabrous-setulose; floral leaves oblong or ovate-lanceolate, 
2.5—4 mm long, 0.5—1.5 mm broad, cuspidate or attenuated acuminate, 
flat, glabrous and smooth along margin. Inflorescence usually almost 
corymbose, consisting of 2-3 fairly dense clusters, with 3-9 flowers 


257 


242 


in each cluster. Flowers almost sessile or on very short, about 0.5 mm 
long, less distinct pedicels. Corolla narrowly infundibuliform or tubular- 
infundibuliform, usually about 5.5 mm long; lobes reflexed, oblong- 
ovate, 2-2.5 mm long, 1—-1.25 mm broad, pink, with large, whitish, 
callous tip, completely glabrous and smooth. Anthers about 1 mm 
long. Ovary and young fruit more or less tuberculate to almost smooth. 
June to August. 

On mountain slopes and meadows in alpine zone.— Caucasus: 
Western Transcaucasia (Abkhazia, Mingerila and Svanetia). Endemic. 
Described from western Abkhazia (Adubulydzh, in valley between 
Mt. Kutysh and Mt. Akhanvash). Type in Leningrad. 

Note. Plants from the alpine meadows of Mt. Anugra in the Gagry 
massif are distinguished from the typical form of the species in having 
a larger, 7-8 mm-long corolla with a slender tube and, probably, belong 
to the separate, more northern geographic race. 


44. A. pontica Boiss. Fl. or. HII (1857) 39; Kreczetowicz in Grossh. 
Fl. Kavk. IV, 27. 

Perennial. Dark green, darkened on drying, subshrub with 
numerous densely foliated sterile shoots, forming small dense mats. 
Flowering stems suberect, mostly 5-10 cm tall, with up to 1-1.5 cm 
long internodes, fairly slender (about 0.5 mm thick), weakly scabrous- 
setulose, furcately branched in upper part or often simple, densely 
foliated. Cauline leaves from linear-oblong (lower) or narrowly linear 
and narrowly linear-lanceolate (upper), 2.5—-12 mm long, 0.4-1 mm 
broad, cuspidate, flat, margin in lower part scabrous-fimbrillate, 
glabrous and smooth in upper part; floral leaves mostly oblong and 
ovate-lanceolate, 2.5-5 mm long, 0.75-1.5 mm broad, more or less 
distinctly narrowed toward base, cuspidate, a few narrowly linear, 
about 0.3-0.4 mm broad, narrowly acuminate. Inflorescence with 1- 
3 clusters of 3-9 flowers each. Flowers almost sessile or on less distinct, 
sometimes, up to 1 mm long, pedicels. Corolla infundibuliform, 5—5.5 
mm long; lobes more or less deflexed, oblong-ovate lobes, about 2 
mm long and 1 mm broad, reddish, with fairly large, whitish, callous 
tip; corolla tube with scattered or isolated fine setae in lower part; 
lobes glabrous and smooth along margin. Anthers about 1 mm long. 
Styles 1.5—2 mm long, free at tip. Ovary finely tuberculate, as in fruit. 
June to August. 

In alpine meadows.— Caucasus: Western Transcaucasia (known 
only from Adzharo-Imeretinsky and Adzharo-Shavshetsky ranges). 
General distribution: Balkans-Asia Minor (eastern part). Described 
from Turkish Armenia. Type in Geneva. 


258 


243 


Series 10. Dolichophyllae V. Krecz. Very low subshrubs with 
abundant densely foliated sterile shoots, forming dense and thick mats. 
Flowering stems 3-7 cm tall, often simple, very densely foliated. 
Cauline leaves narrowly linear, up to 12-18 mm long, with margins 
more or less revolute, all considerably exceeding internodes (something 
not seen in representatives of other series known to us); floral leaves 
linear- or oblong-ovate-lanceolate, 3—7.5 mm long, exceeding or almost 
equaling corolla of flowers (in all other species we know, they are 
considerably shorter than flowers). Flowers almost sessile, 3—9 together 
in dense capitate clusters, enveloped by floral leaves. Corolla tubular- 
infundibuliform, 5-6.5 mm long; lobes oblong-ovate or ovate- 
lanceolate, considerably shorter than tube, pink, with very small whitish 
thickened tip, glabrous and smooth. Anthers linear, about 1.5 mm 
long (longer than in other species). Styles 1-2 mm long. Fruit covered 
by cuneate subacute tubercles. 

Iranian high-mountain species reaching southern Transcaucasia. 


45. A. affinis Boiss. et Huet, Diagn. ser. II, 2 (1856) 110; Krecz. 
in Grossh. Fl. Kavk. IV, 27.—A. cynanchica var. y. affinis Boiss. FI. 
or. III (1857) 41, quoad plantam armeniacam. 

Perennial. Grayish-green subshrub with numerous densely foliated 
sterile shoots, forming thick, dense mats. Flowering stems suberect or 
ascending, 3—7 cm tall, with up to 0.5—1 cm long internodes, not very 
slender (0.5—0.75 mm thick), weakly scabrous-setulose in upper part, 
toward apex weakly scabrous or subglabrous, single or apex furcately 
branched. Cauline leaves narrowly linear, 3—12(15) mm long, 0.25— 
0.75 mm broad, mucronate, with somewhat revolute margin, weakly 
scabrous-setulose along margin, mainly in lower part; floral leaves 
narrowly linear-lanceolate, 3-7 mm long, 0.5—1 mm broad, cuspidate, 
or narrowly acuminate, very weakly scabrous-setulose to almost smooth 
along margin, sometimes mixed with rudimentary, almost filiform 
leaves. Flowers almost sessile or on short less distinct pedicels, 3—7 
together in capitate clusters at end of stem or branches. Obvolute 
envelope of floral leaves exceeding corolla. Corolla narrowly 
infundibuliform, 5—5.5 mm long; lobes more or less deflexed, oblong- 
ovate, 2.25-2.5 mm long, about 0.75 mm broad, pink, with small, 
whitish, callous tip, entirely glabrous and smooth. Anthers linear, about 
1.5 mm long. Ovary and young fruits covered by elongated, cuneate 
tubercles. June to August. 

In rocks of subalpine zone.— Caucasus: Southern Transcaucasia 
(along boundary with Turkey). General distribution: Armenia- 
Kurdistan. Described from vicinity of Erzerum. Type in Geneva. 


259 


244 


46. A. dolichophylla Klok. sp. n. in Addenda XXII, 706. 

- Perennial. Glaucescent-green, comparatively pale colored subshrub 
with numerous, densely foliated, sterile shoots, forming fairly dense 
mats. Flowering stems ascending, 3-5 cm tall, with up to 1.5 cm long 
internodes, not very slender (about 0.75 mm thick), scabrous-setulose 
throughout; simple or furcately branched at tip, with more or less 
elongated, densely foliated branches; lower leaves oblong or linear- 
lanceolate, 1.5—3 mm long, about 0.75 mm broad (at base), more or 
less acuminate, flat, almost smooth along margin; cauline leaves 
narrowly linear, 7-18 mm long, 0.5—1 mm broad, cuspidate, with 
revolute margin, fairly weakly scabrous-setulose along margin; lower 
leaves densely aggregated, upper sparse, but also longer than 
internodes; floral leaves from linear to oblong-ovate-lanceolate, 3—7.5 
mm long; 0.5-1.5 mm broad (longest of these slightly shorter than 
flowers), thinly tapering, flat, weakly scabrous-setulose along margin. 
Flowers sessile or subsessile, 5-9 together at ends of stems or their 
branches in fairly dense, capitate clusters; obvolute envelope of floral 
leaves almost equaling corolla of flowers. Corolla tubular- 
infundibuliform, 5.5—6.5 mm long; lobes more or less reflexed, oblong- 
lanceolate or oblong-ovate-lanceolate, mucronate, 2-3 mm long, about 
1 mm broad, pink, with small whitish callous tip, glabrous and smooth. 
Anthers 1.25—1.5 mm long, linear. Style 1.5—2.5 mm long. Ovary and 
young fruits covered with cuneate subacute tubercles. June to August. 

In rocks in alpine and subalpine zone.—Caucasus: Southern 
Transcaucasia (probably occurs along boundary with Turkey, precise 
localities not yet established). General distribution: Armenia-Kurdistan. 
Described from Kars district (mountains above village of Bazat). Type 
in Leningrad. 


Series 11. Pseudograveolentes V. Krecz.—Weakly woody 
subshrubs, not forming a mat. Flowering stems 20—40 cm tall, slender, 
with elongated upper internodes. Middle cauline leaves narrowly linear 
to almost setaceous, up to 2.5 cm long, with margins strongly revolute; 
floral leaves oblong-lanceolate, very small. Inflorescence spreading, 
paniculate. Flowers on very short or more distinct pedicels. Corolla 
very small or large (up to 3.5-4.mm long); lobes obtusely acuminate, 
one and a half to two times as short as tube, setulose or glabrous and 
smooth, pink, with small whitish, callous tips. 

Transcaucasian species of lower mountain belt. 

Note. This series was included by Kreczetowicz in the cycle 
Parviflorae V. Krecz. and placed second after series Leptocladae V. 
Krecz., which includes A. rumelica Boiss. In most respects it really 


260 


245 


resembles our series Rumelicae Klok., but apparently has arisen 
independently of the latter series. 


47. A. woronowii V. Krecz. in Grossh. Fl. Kavk. IV (1934) 25. 

Perennial. Weakly woody subshrub, sometimes forming very lax 
mats. Flowering stems suberect to almost erect, 20-40 cm long, with 
up to 3—4.5 cm long internodes, slender (0.5—0.75 mm thick), fairly 
weakly scabrous-setulose in lower part, glabrous and smooth upward, 
furcately or ternately branched above middle, with very slender 
branches, sparsely foliated; middle cauline leaves narrowly linear to 
almost setaceous, 10-25 mm long, 0.5-0.75 mm broad, cuspidate, 
flat; floral leaves oblong-ovate-lanceolate, 1-2 mm long, 0.25-0.75 
mm broad, tapering acuminate, along margin (just as upper cauline 
leaves) glabrous and smooth or nearly so. Inflorescence spreading, 
paniculate. Flowers on ends of branches, 2—3 together, rarely S-7. 
Pedicels very short or rarely more distinct, 1-2 mm long. Corolla 
infundibuliform, 2.5—3.5 mm long; lobes weakly deflexed, oblong- 
ovate-lanceolate, mucronate, 1—-1.5 mm long, 0.5—0.75 mm broad, pink, 
with small whitish, callous tip, uniformly setulose. Anthers about 0.75 
mm. Styles 1—1.5 mm long, free at tip. Fruits about 2 mm long, finely 
tuberculate. June to July. 

On stony mountain slopes.— Caucasus: Southern Transcaucasia 
(near Artvin district). General distribution: Armenia-Kurdistan. 
Described from environs of Artvin (Lomashen). 


48. A. kemulariae Manden. in Zap. po. sist. i geogr. rast. bot. 
inst. AN GruzSSR, 17 (1953) 58; Manden. in Fl. Gruzii, VIII, 14.— 
Ic.: Fl. Gruzii, VIII, tab. 360. 

Perennial. Grayish-green subshrub. Flowering stems suberect, about 
30-35 cm tall, with up to 5—5.5 mm long internodes, slender, scabrous- 
setulose in lower part, glabrous and smooth in upper part, divaricately 
branched above middle, sparsely foliated. Basal leaves oblong, about 
3 mm long and 0.75 mm broad, acuminate; cauline leaves narrowly 
linear to almost setaceous, 3-23 mm long, 0.2—-0.3 mm broad, cuspidate 
or attenuated into very short apical mucro, with revolute margin, more 
or less deflexed from stem; lower leaves weakly scabrous-setulose 
along margin, upper ones glabrous and smooth; floral leaves linear to 
oblong-lanceolate, 1—2.5 mm long, 0.2-0.5 mm broad, thinly acuminate. 
Inflorescence paniculate, spreading. Flowers on terminal branches, 
grouped in threes. Pedicels very short. Corolla tubular-infundibuliform, 
44.5 mm long; lobes deflexed, oblong-ovate-lanceolate, almost 2 times 
as short as tube, pink with whitish callous tip, glabrous and smooth. 
Ovary and fruit finely tuberculate. June to August. 


261 


246 


On rocks in lower mountain belt.—Caucasus: Western 
Transcaucasia. Endemic. Described from western Georgia (Lechkhumsk 
District, estuary of Ladzhanura River, Orpiri). Type in Tbilisi, cotype 
in Leningrad. 


Series 12. Graveolentes Klok.—Subshrub, with somewhat 
developed or, almost absent, sterile shoots, sometimes forming very 
lax mats. Flowering stems 10—60 cm tall, virgate. Middle cauline leaves 
linear-subulate, up to 20-40 mm long, thick (as all others), with 
revolute margin; floral leaves mostly oblong-lanceolate, very small. 
Inflorescence paniculate but narrow. Flowers on more or less distinct 
pedicels. Corolla infundibuliform or tubular-infundibuliform, 2.5—6.5 
mm long; lobes obtuse or mucronate, almost equaling tube or (rarely) 
considerably shorter, pink, usually subglabrous. Fruits comparatively 
large, up to 3—4.5 mm long, densely set with flat, obtuse or acuminate, 
scaly outgrowths, rarely almost without scales (glabrate). 

To this series belong the steppe species growing exclusively on 
sands. In the system of Kreczetowicz they are classified in the cycle 
Lepidophorae V. Krecz., without further division into series. Besides 
our species, it also includes A. littoralis Sibth. et Sm. from Asia Minor. 
Series Graveolentes Klok. sharply differs from all other series and 
can be regarded as a separate section of the subgenus, for which it is 
possible to retain the name proposed by Kreczetowicz for the cycle. 


49. A. setulosa Boiss. Diagn. ser. I, 10 (1849) 61; FI. or. III, 40 


sub A. littoralis in textu; Grecescu, Consp. fl. Rom. 266; Prodan, FI. 


pl. cresc. Rom. I, 967; Chrshan. in Vizn. roslin URSR, 309.—Exs.: 
Lang and Szovits, Herb. ruth. cent. II, No. 108. 

Perennial. Grayish-green subshrub with weakly developed sterile 
shoots, woody, thick primary branchings of axis 1.5—4 mm in diameter. 
Flowering stems numerous, suberect, 30-50 cm tall, shortly setose- 
hairy in lower part together with leaves, glabrous and smooth or weakly 
scabrous upward, together with leaves, branched above or sometimes 
below middle, densely foliated at base, with sterile shoots in axils of 
basal leaves. Basal leaves from ovate, obtuse to oblong, short- 
acuminate, 1.5 mm long, 0.75—1 mm broad; middle cauline leaves 
linear-subulate, 9-30 mm long, 0.5—1.25 mm broad, cuspidate, with 
revolute margin and prominent vein beneath, thicker, somewhat fleshy; 
upper leaves (in inflorescence) oblong-lanceolate, cuspidate, 2-5 mm 
long, 0.5-0.75 mm broad; floral leaves similar in shape, 1.5-2.25 mm 
long, 0.3-0.5 mm broad. Inflorescence paniculate, not broad. Flowers 
on terminal branches in groups of three, middle flower of cyme almost 
sessile; lateral flowers on more or less distinct pedicels. Corolla 


262 


247 


infundibuliform, 4—4.5 mm long; lobes oblong-elliptic or ovate, 
somewhat narrowed toward base, obtuse, scattered setulose along 
margin, 2.25—2.5 mm long, 1—1.25 mm broad, somewhat longer than 
tube. Styles 1-2 mm long, free at tip. Fruits 2.5—-4 mm long, densely 
covered with mucronate small scales. July to August. 

On littoral sands.— European USSR: Bessarabia (southern part), 
Lower Dniéper (southwestern part). General distribution: Balkans 
-Asia Minor (Black Sea coast of Romania). Described from vicinity of 
Odessa. Type in Geneva, cotypes in many places (Leningrad, Kiev). 

Note. A littoral endemic species of the northwestern Black Sea 
coast, extending from Constantsa in Romania to the Perekop Isthmus. 
Not seen on the coast of the Sea of Azov. On the sands of the Lower 
Dnieper there is a widespread form not always clearly distinguishable 
from A. graveolens M.B., but evidently belonging to this littoral race. 


50. A. savranica Klok. sp. n. in Addenda, XXII, 707.—A. 
graveolens auct. fl. ucr. pro minima parte, non M.B. 

Perennial. Dark green subshrub. Flowering stems obliquely 
ascending, 12-25 cm tall, along with leaves scabrous-setulose to the 
very top (setae in lower part of stems conspicuous and dense, toward 
top not so dense, but distinct), branched below middle and upward, 
densely foliated, with sterile shoots in axils of lower leaves. Basal 
leaves from oblong-ovate (obovate) to linear-oblong, 1-3 mm long, 
about 0.5 mm broad, short-acuminate or cuspidate; middle cauline 
leaves linear-subulate, 3-16 mm long, 0.5-0.75 mm broad, cuspidate 
with revolute margin, thicker, upper leaves in inflorescence oblong- 
lanceolate, up to 4.5 mm long and 1 mm broad; floral leaves in 
inflorescence similar in shape, 1.5-2.5 mm long, 0.5—1 mm broad, 
acuminate. Flowers almost sessile or on short pedicels. Corolla 
infundibuliform, 3—3.5 mm long; lobes more or less deflexed, oblong, 
obtuse, 1.5-1.75 mm long, about 0.75 mm broad, approximately 
equaling tube. Style about 1 mm free or more or less sessile. Fruits 
1.75—3 mm long, fairly densely covered with acute small scales. June 
to July (Plate XIII, fig. 3). 

Alluvial sands.— European USSR: Middle Dnieper (above Southern 
Bug, Kodyma and Savran). Endemic. Described from Odessa region. 
Type in Leningrad. 


51. A. graveolens M.B. sec. Bess. in Roem. et Schult. Mant. ad 
Syst. veg. III (1827) 376; Ldb. Fl. Ross. II, 399.—A. pseudograveolens 
M. Pop. et Chrshan. in Byull. Mosk. Obshch. ispyt. prir. L, 5—6 (1945) 
94.—Exs.: GRF, No. 1068. 


263 


248 


Perennial. Bright green subshrub, with more or less, developed 
sterile shoots, sometimes forming mats. Flowering stems obliquely 
ascending, 10-35 cm tall, thicker scabrous-setulose though not densely 
in lower part, glabrous and smooth in upper part, branched from middle 
or below, fairly densely foliated, with sterile twigs in axils of lower 
leaves, sometimes also with elongated sterile shoots. branching out 
from primary branches of axis. Basal leaves from oblong to oblong- 
lanceolate, 2-5 mm long, about 1 mm broad; middle cauline leaves 
narrowly linear-subulate, up to 15-40 mm long, 0.5—1.25 mm broad; 
cuspidate, with revolute margin, thicker, those situated below more or 
less scabrous, remaining glabrous and smooth; floral leaves oblong- 
lanceolate, 1.5—3 mm long, about 0.5 mm broad, smooth. Inflorescence 
(dichasially) paniculate. Flowers terminal on branches, in groups of 
(twos), threes, rarely solitary, almost sessile or on up to 1-1.5 mm 
long pedicels. Corolla infundibuliform, 3.5—4.5 mm long; lobes 
deflexed, oblong, obtuse, diffusely setulose along margin, 1.75—2 mm 
long, 0.75—1 mm broad, almost equaling tube, pale pink; corolla with 
exception of lobes subglabrous. Styles 1-2 mm long, free above middle 
or almost from base. Fruits 3—4 mm long, more or less densely covered 
by small obtuse or subobtuse scales, very rarely almost without scales 
(f. pseudoleiograveolens Klok.). June to August. 

Alluvial sands.—European USSR: Middle Dnieper (east of 
Dnieper), Volga-Don (in Don Basin), Lower Don (also in Don Basin). 
Reports for other districts not verified or doubtful. Endemic. Described 
from Don Region (village of Ellanskaya). Type in Leningrad, cotype 
in Kiev. 


52. A. leiograveolens M. Pop. and Chrshan. in Byull. Mosk. 
obshch. ispyt. prir. otd. biol. L, 5-6 (1945) 96; Chrzhanovsk. in Vizn. 
roslin URSR (1950) 309.—Exs.: Gerb. fl. URSR, No. 94 (sub A. 
graveolens M.B.). 

Perennial. Bright green, weakly woody subshrub with slender 
primary branchings of axis (up to 1 mm in diameter), often forming 
lax mats. Flowering stems obliquely ascending, 10-30 cm tall, thicker 
than in A. graveolens M.B., weakly scabrous-setulose at base, above 
glabrous and smooth, more or less branched, fairly densely foliated, 
with sterile twigs in axils of lower leaves and often with elongated 
sterile shoots, emerging from primary branches of axis. Basal leaves 
linear-oblong, very small, badly preserved (about 2 mm long, and 0.3 
mm broad); middle cauline leaves narrowly linear-subulate to almost 
filiform, 10-25 mm long, 0.3-0.75 mm broad, cuspidate, with revolute 
margin, glabrous and smooth; upper leaves (in inflorescence) oblong- 
or linear-lanceolate, up to 5 mm long, and 1 mm broad; floral leaves 


264 


249 


similar in shape, usually 2—2.5 mm long and about 0.5 mm broad 
(sometimes up to 4 mm long and about 0.3 mm broad), smooth. 
Inflorescence paniculate. Flowers almost sessile or on up to 2 mm 
long pedicels. Corolla infundibuliform, 3.5-3.75 mm long, lobes 
oblong, obtuse, 1.75-2 mm long, 0.7-1 mm broad, a bit longer than 
tube, pale pink, glabrous and smooth. Styles 1.2-1.75 mm long, free 
at tip. Fruits about 3 mm long, scales almost absent. June to August. 

Alluvial sands.— European USSR: Middle Dnieper (along Dnieper 
in Kiev and Cherkessia regions, UkrSSR). Endemic. Described from 
sands of Middle Dnieper. Type in Kiev, cotype in Leningrad. 


53. A. diminuta Klok. sp. n. in Addenda XXII, 707.—A. 
danilewskiana Woron. et Schelk. in sched. herb. Fl. cauc. VII, No. 
347; Tr. Tifl. bot. sada, XIX, 69, non Basin.; Maiorov, E’ol. pust. u 
podnozh. Dag. 101, tabl. 21.—Exs.: Herb. Fl. Cauc. No. 347 (sub A. 
danilewskiana Basin.). 

Perennial. Glaucescent-green subshrub with elongated (but not 
very long) sterile shoots, branching out from comparatively thick 
primary branches of axis. Flowering stems usually fairly numerous, 
ascending or suberect, 15-35 cm tall, glabrous and smooth from base 
and throughout, more or less branched above middle, with shortened 
sterile axillary twigs. Basal leaves from broadly deltoid-ovate to ovate- 
oblong, 1—4 mm long, 0.75-1.5 mm broad, obtuse or subobtuse; middle 
cauline leaves linear-subulate, 6-20 mm long, 0.5—1.25 mm broad, 
cuspidate, with revolute margin, very weakly or indistinctly scabrous 
along margin, mostly considerably shorter than internodes, thicker; 
floral leaves oblong-ovate-lanceolate, 1.2-1.5 mm long, about 0.75 
mm broad, smooth. Inflorescence paniculate, not many-flowered. 
Flowers on terminal branches gathered in (twos) threes, almost sessile 
or on about 1 mm long thin pedicels. Corolla tubular-infundibuliform, 
3.54.5 mm long; lobes deflexed oblong or oblong-ovate, somewhat 
obtuse, 1.5-2.5 mm long, about 1 mm broad, almost equaling tube, 
pale pink, completely glabrous and smooth. Styles about 1 mm long, 
more or less conferruminate. June to July. 

On sands.— European USSR: Lower Volga (southwestern part). 
Caucasus: Dagestan (northern part). Endemic. Described from Dagestan 
(Kumtorkali). Type in Leningrad. 


54. A. laevissima Klok. sp. n. in Addenda XXII, 708.—A. 
graveolens Krecz. in Fl. Yugo-Vost., VI (1936) 257, non M.B. 

Perennial. Dark green subshrub, often with elongated sterile shoots, 
branching out from primary branches of axis. Flowering stems 
ascending or suberect, 35-60 cm tall, glabrous and completely smooth 


265 


250 


from base throughout, branched from middle or above, lower branches 
with shortened sterile axillary twigs. Basal leaves badly preserved, 
middle cauline leaves linear-subulate, 10-40 mm long, 0.5—1 mm broad, 
cuspidate, with revolute margin, smooth along margin, thicker and 
somewhat fleshy, often more or less curved; floral leaves oblong- 
lanceolate, very small, 1.5-1.8 mm long, 0.3-0.5 mm _ broad. 
Inflorescence paniculate (more condensed, but less dense than in A. 
graveolens M.B.). Flowers on terminal branches gathered in threes, 
middle flowers or cyme usually almost sessile, lateral on short, up to 
1.5 mm long, pedicels. Corolla tubular-infundibuliform, 4.5-6 mm 
long; lobes weakly deflexed, oblong-ovate or oblong-ovate-lanceolate, 
obtuse or subobtuse, 1.5—2.25 mm long, about 0.75 mm broad, almost 
2 times as short as tube. Styles 2-2.5 mm long, free at tip or from 
middle. Fruits 3-4.5 mm long, densely covered with short ligulate 
scales, ripe fruit reddish brown. June to July (Plate XIII, fig. 1). 

Alluvial sands.— European USSR: Volga-Don (southern part), 
Lower Don (northern part), Volga Region (southern part), Lower Volga 
(along River Ural). Endemic. Described from Boguchar district of 
Voronezh Region of RSFSR. Type in Leningrad. 

Note. The species described above clearly occupies, on the basis 
of its morphological characteristics and geographical distribution, an 
intermediate position between A. graveolens M.B. and A. danilewskiana 
Basin. Often, it has been misidentified as the former-species, rarely as 
the latter species, to which, apparently, it is closer. 


55. A. danilewskiana Basin. in Bull. phys.-math. Acad. Sc. Pétersb. 
II (1844) 202; Boiss. Fl. or. III, 40; Korsh. Tent. 190; B. and O. 
Fedtsch. Consp. Fl. Turk, 135; Kreczetowicz in Fl. Yugo.-Vost. VI, 
258.. 

Perennial. Glaucescent-pale green subshrub, with weakly developed 
sterile shoots. Flowering stems rising upward to suberect, 30-60 cm 
tall, with up to 4-6 cm long internodes, more or less arched, obtusely 
4-angled to almost cylindrical in lower part, 1-2 mm in diameter, very 
weakly scabrous-setulose or scabrous, completely smooth above, 
furcately (doubly furcately) and ternately branched from middle or 
below, often almost from base, sparsely foliated. Basal leaves 
squamiform, more or less crowded, from ovate-deltoid to oblong-ovate, 
1.5-3 mm long, 0.75—1 mm broad, narrowed toward apex, obtuse or 
mucronate, scabrous along margin; middle cauline leaves separated, 
narrowly linear, 7-24 mm long, 0.75—-1 mm broad, cuspidate, with 
more or less revolute margin, scabrous along margin; at base joined 
in a noticeable constriction (0.5—1 mm high, or more), more or less 
patent and curved in arch; upper leaves (in inflorescence) linear- or 


266 


251 


oblong-lanceolate, 2.5-10 mm long, 0.75-1.5 mm broad, fairly 
narrowly acuminate, flat, along margin, weakly scabrous or strongly 
fimbrillate; floral leaves oblong-lanceolate, 1.5—4 mm long, 0.5—1 mm 
broad, acuminate, scabrous along margin. Inflorescence paniculate, 
elongated but comparatively narrow. Flowers 1—3 together on terminal 
branches of inflorescence, almost sessile or on noticeable, up to 4—5 
mm long pedicels. Corolla tubular-infundibuliform, 5.5—6.5 mm long; 
lobes reflexed, oblong-lanceolate, somewhat narrowed toward base, 
mucronate, 1.5—2 mm long, 0.75-1 mm broad, pale pink, with less 
distinct whitish, callous tip. Corolla tube elongated, slender, cylindrical 
in lower part, glabrous and smooth, lobes smooth along margin, nearly 
2 times as short as tube. Anthers about 1.5 mm long. Styles 1.5—2 mm 
long, unequal, free at tip. Fruits 2.5—4 mm long, subglobose, densely 
covered by white, lanceolate, obtusely acuminate scales, beneath brown. 
May to August. 

On sands.—European USSR: Lower Volga (along left banks of 
Ural River). Western Siberia: Upper Tobol (southwestern part). Soviet 
Central Asia: Aralo-Caspian. Endemic. Described from northern tip of 
Ust-Urt. Type in Leningrad. 

Note. This species, probably along with the former, should be 
placed in the separate series Danilewskianae Klok. 


Section 5. GALIoIwEAE DC. Prodr. IV (1830) 585, p.p.; sect. 
Cynanchica DC. § Galioideae Boiss. Fl. or. III (1875) 27, p.p.; 
Schumann in Pflanzenfam. IV, Abt. 4, 5, 149.—Corolla shortly 
campanulate, white. Stigma globose. Leaves filiform, linear, lanceolate 
or ovate-oblong. Stem weakly branched or unbranched. 


Series 1. Octonariae Klok.—Stems glabrous or pubescent with 
white long hairs. Leaves 6-8 in a whorl, linear or filiform. Corolla 
lobes with short, thick incurved appendage. 


56. A. octonaria Klok. in Bot. mat. Gerb. Bot. inst. AN SSSR, 
XVIII (1957) 229.—A. galioides M.B. Fl. taur.-cauc. III (1819) 104, 
p-p.; DC. Prodr. IV, 585, p.p.; Ldb. Fl. Ross. II, 399, p.p.—A. glauca 
auct. non Besser; Schmalh. Fl. II (1897), 10, p.p.; Grossh. Fl. Kavk. 
IV, 24, p.p.—Exs.: GRF, No. 719 (sub A. glauca Bess.); Herb. FI. 
Ucr. No. 95. 

Perennial. Rootstock thick, short, woody, dark brown. Stems 
numerous, straight, sterile, short, densely foliated; flowering stems 
20-80 cm long, branched only in inflorescence, terete in lower part, 
above indistinctly 4-angled, glabrous, lustrous. Leaves in whorls of 8, 
filiform, (20)25(45) mm long, (0.25)0.5(0.75) mm broad, with highly 


267 


252 


revolute margin, with short thick cusp, sessile, glabrous or diffusely 
setulose, gray-blue, paler beneath; floral leaves 4-5 in number, 3-4 
times as short as stems, long acuminate, uppermost leaves elliptic or 
lanceolate. Inflorescence spreading, with long lateral branches, 
paniculate-corymbose. Peduncles and pedicels glabrous; pedicels short, 
clavate below flowers, equaling flowers or shorter; peduncles bearing 
cymes, doubly ternately branched, cymes 5-6 flowered. Corolla 
campanulate, 3-3.5 mm across, with broad, short tube, equaling lobes 
or shorter; lobes oblong-ovate, tapered into thick short incurved tip. 
Stamens with filaments, equaling anthers, inserted somewhat below 
upper margin of tube. Style bifid at 2/3 length; stigmas globose. Fruits 
binate, 2 mm long, 3 mm broad, granulate, rugose. Flowering May to 
June. Fruiting July to August (Plate XIV, fig. 3). 

Steppes— European USSR: Upper Volga, Volga-Kama, Upper 
Dnieper, Middle Dnieper, Volga-Don, Volga Region, Upper Dniester, 
Bessarabia, Black Sea Region, Crimea (only steppe part), Lower Don, 
Lower Volga. Caucasus: Ciscaucasia. Soviet Central Asia: Aralo- 
Caspian, Balkhash Region. General distribution: Balkans-Asia Minor 
(northern ‘steppe part of Balkan Peninsula). Described from environs 
of Taganrog. Type in Leningrad. 


57. A. campanulata (Vill.) Klok. in Bot. mat. Gerb. Bot. inst. 
AN, XVIII (1957) 226.—Galium campanulatum Villars. Dauph. II 
(1787) 326, tab. VII; Roem. and Schult. Mantissa, HI, 174.—G. 
glaucum auct. non L.: Roem. and Schult. Syst. veg. III (1818) 210, 
p.p.—Asperula glauca Bess. Suppl. III ad Cat. pl. (1814) 4 and Enum. 
pl. Volh. (1822) 7; Schmalh. Fl. II, 10, p.p.; Grossh. Fl. Kavk. IV, 24 
p.p.—G. galioides auct. non M.B.: DC. Prodr. IV (1830) 585, p.p.; 
Ldb. FI. Ross. II, 399; Boiss. Fl. or. III, 44, p.p.—Ic.: Hegi, Ill. Fl. 
VI, 1, tab. 247, 4a.—Exs.: Fl. Pol. exs. No. 44 (sub A. galioides 
M.B.); Fl. Gall. et Germ., exs. No. 79. 

Perennial. Rootstock thick, woody, brown. Stems several, straight, 
60 cm tall, branched, thickened at nodes, lustrous, glabrous. Leaves in 
whorls of 8, upper ones and on branches in whorls of 6, linear, 15— 
40 mm long, 0.5—1 mm broad, with short cusp, easily revolute and 
here hardly scabrous along margin, green on both surfaces, glabrous; 
floral leaves 1-2-4, very small, 1.5—2 mm long, 0.5—0.75 mm broad, 
lanceolate or elliptic. Inflorescence paniculate, very lax. Peduncles 
doubly-triply furcately or ternately branched, glabrous together with 
pedicels; cymes 5-9 flowered; pedicels short, equaling flowers or 
shorter, clavate below flowers. Corolla white, campanulate, 1-2 mm 
across with broad, short tube, shorter than lobes; lobes oblong with 
involute tip. Stamens with large, elliptic anthers, shorter than filaments. 


268 


253 


Style short, bifid almost from base, with hypogynous disc at base. 
Fruits often with one mericarp, almost globose, 1.5 mm long and 
broad, rugose, surface glabrous when ripe. June to July. 

Dry mountain slopes.— European USSR: Middle Dnieper, Upper 
Dniester. Described from France. General distribution: Atlantic Europe, 
Balkans. Type unknown. 


58. A. tyraica Bess. Enum. pl. Volh. (1822) 41; Stev. in Buil. 
Soc. Nat. Mosc. XXIX, 362; Pachossk. Nov. dan. o fl. Khersonsk. 
gub. 13 and in Tr. Bess. obshch. est. III, 14.—A. galioides B. tyraica 
DC. Prodr. IV (1830) 585; Ldb. Fl. Ross. II, 400; Schmalh. FI. II, 10; 
Grossh. Fl. Kavk. IV, 24. 

Perennial. Rootstock thin, horizontal, dark brown. Stems one or 
few, 30-60 cm tall, straight or ascending, branched, glabrous, lustrous, 
more or less pubescent with white upright hairs on lower internodes, 
especially dense at its nodes. Leaves in whorls of 8, upper ones and 
those in branches in whorls of 6, linear, narrow, (20)35(40) mm long, 
0.5—1 mm broad, upper ones erect, apex with thin often curved cusp, 
revolute along margin and here scabrous from scattered setae; lower 
leaves often with soft, patent pubescence; floral leaves in whorls of 1- 
2-6, minute, 2-3 mm long, 0.5—1 mm broad, glabrous. Inflorescence 
paniculate, with long lateral branches. Peduncies doubly-triply 
branched, glabrous, cymes 5-9 flowered; pedicels short, equaling 
flowers or somewhat shorter. Corolla white, 3—-3.5 mm across, 
campanulate with broad, short tube, shorter than lobes; lobes ovate, 
with thick incurved tip. Anthers elliptical on longer filaments. Style 
bifid from middle, with hypogymous disc at base. Fruit often with one 
oblong mericarp, 2 mm long, 0.75 mm broad, glabrous, rugose, 
granulate. June to August. 

On calcareous mountain slopes, virgin steppe and along borders of 
fields.— European USSR: Middle Dnieper, Upper Dniester, Bessarabia. 
Described from Podolia. Type in Leningrad. 

Note. The species is close to A. campanulata (Vill.) Klok., but 
intermediates between these species are absent, as noted by I. 
Pachossky, who observed it in the wild. 


59. A. galioides M.B. Fl. taur.-cauc. I (1808) 101; DC. Prodr. IV, 
585, p.p.; Ldb. Fl. Ross. II, 399, p.p.; Boiss. Fl. or. III, 44, p.p.— A. 
glauca auct. non Bess.: Schmalh. FI. II, 10, p.p.; Grossh. Fl. IV, 24, 
p-p. 

Perennial. Rootstock slender, branched, dark brown. Stems from 
one to several, 15-50 cm tall, sterile, short, densely foliated; flowering 
stems straight, terete, indistinctly 4-angled upward, branched, glabrous, 


271 


254 


lustrous. Leaves in whorls of 8, upper leaves in whorls of 6, linear, 
(7)15(25) mm long, (0.25)1(1.5) mm broad, with short cusp at apex, 
revolute along margin and here sharply scabrous, elsewhere glabrous 
or above scabrous from fine, upward directed setae, paler beneath 
with one, sharply prominent vein, sessile; floral leaves paired or 
solitary, in basal axes in whorls of 4—6, elliptic, acute, 1-2 mm long, 
0.5-0.75 mm broad, glabrous together with peduncles and pedicels; 
pedicels clavate below flowers. Inflorescence paniculate or corymbose- 
paniculate, lax. Peduncles doubly branched, cymes 2-6 flowered. 
Flowers on short pedicels, equaling flower. Corolla 3.5 mm across, 
white, campanulate, with broad, short tube, equaling lobes or somewhat 
shorter or slightly longer; lobes ovate, acute, glabrous. Stamens inserted 
below upper margin of tube, filaments longer with elliptic anthers. 
Styles short, included in tube, bifid in upper half, with globose stigmas. 
Fruits binate or with one mericarp, 2 mm long, 2.5 mm broad; mericarps 
oblong, glabrous, rugose. June to September. 

On stony mountain slopes, screes.—European USSR: Crimea (only 
mountainous part). Caucasus: Ciscaucasia. Endemic. Described from 
Crimea. Type unknown. 


60. A. xerotica Klok. in Bot. mat. Gerb. bot. inst. AN SSSR, 
XVIII (1957) 228. 

Perennial. Rootstock slender, branched, reddish dark brown. Stems 
numerous, 15—40 cm tall, predominantly sterile, densely foliated; 
flowering stems 6-9, straight or in upper part branched, with short 
erect branches, almost terete in lower part, 4-angled upward, densely 
and softly pubescent only in lower part or throughout. Leaves in whorls 
of 6-8, linear, narrow, (17)18(23) mm long, 0.25—0.5 mm broad, 
cuspidate, with strongly revolute margin, strongly scabrous on both 
surfaces, especially lower leaves; upper leaves glabrous above, scabrous 
beneath; floral leaves in whorls of 4, long acuminate, uppermost leaves 
solitary, elliptic, very small, 2-4 mm long, 0.25-0.5 mm broad, 
glabrous together with peduncles and pedicels. Pedicels clavate under 
flowers, shorter than flowers. Inflorescence not long, more or less 
compact with short, upward directed branches; cymes ternately 
branched, 4—6-flowered. Corolla white, 3.5 mm across, campanulate, 
with broad, short tube, equaling lobes or somewhat shorter, or slightly 
longer; lobes ovate, with incurved tip. Stamens inserted below lobes. 
Anthers elliptic, browning on drying. Style bifid in upper third. Ripe 
fruits unknown. Flowering May. 

On stony steppes, rocks.— European USSR: Crimea. Endemic. 
Described from Tarkhankut Peninsula. Type in Leningrad. 


255 


Plate XV. 
1. Asperula oppositifolia Rgl. and Schmalh., upper and lower parts of plant, 
fruit.—2. A. laevis Schischk., general appearance of plant, unfolded corolla.— 
3. Callipeltis cucullaris (Jusl.) Rothm., general appearance, flower with 
bracteoles.— 4. Leptunis trichoides (J. Gaj.) Schischk., general appearance, 
unfolded corolla, fruit. 


269 


272 


256 


Series 2. Paniculatae Pobed.—Stems glabrous. Leaves in whorls 
of 4—6, lanceolate, ovate-lanceolate, ovate-oblong, elliptic or linear. 
Lobes of corolla obtuse or rounded at tip. 


61. A. paniculata! Bge. in Ldb. FI. alt. I (1829) 140; Ldb. FI. 
Ross. II, 400; Kryl. Fl. Zap. Sib. X, 2570.—Ic.: Ldb. Ic. pl. Fl. Ross. 
IV, tab. 301. 

Perennial. Rootstock funiform, branched, about 1 mm thick. Stems 
few in number, simple or branched, straight, 4-angled, glabrous, 15— 
50 cm tall. Lower leaves in whorls of 4, remaining in whorls of 6, 
leaves stiff, lanceolate or ovate-lanceolate, acute or short acuminate, 
with short, white setae along margin, dark green and more or less 
lustrous above, paler beneath, 1.5—3(5) cm long, and 3-8 mm broad. 
Flowers aggregated at end of stems and branches in lax panicle. Corolla 
broadly campanulate, whitish (brownish or drying), 4-fid up to middle, 
about 3 mm long. Mericarps smooth, almost globose, about 2 mm in 
diameter. June to July (Plate XIV, fig. 1). 

On stony and grassy slopes, rocks and shingles. In mid-montane 
belt.—Western Siberia: Altai (often), Angara-Sayan (along Yenisei 
tributaries): Golubaya, Kazyr-Suk rivers). General distribution: Hsin-- 
hsiang (Borgaty Creek). Described from Altai. Type in Leningrad. 

Note. The disjunct geographic distribution of this species is 
noteworthy. Widely distributed in the Altai, A. paniculata Bge. was 
collected in the rocks along the Yenisei River (Angara-Sayan) and 
discovered by Regel in the Chinese province of Hsin-hsiang in the 
Dzhungarian Ala Tau; this species has not been found in the Saur and 
Tarbagatai ranges and the Dzhungarian Ala Tau within the USSR. 


62. A. kryloviana Serg. in Trav. de I’Inst. Sc. de Biologie, Tomsk, 
II (1936) 69; Kryl. Fl. Zap. Sib. X, 2571. 

Perennial. Rootstock funiform, about 1 mm thick. Stems straight, 
4-angled, smooth and glabrous, branched almost from base, 15—20 cm 
tall. Leaves in whorls of 4—6, lower leaves ovate; upper leaves stiff, 
more or less lustrous above, dull with one vein beneath, along margin 
scabrous and clingy because of fine spines, 1-1.5 cm long, and 2-5 
mm broad. Peduncles 3 in number, rarely 2, emerging from axils of 
lanceolate or almost linear floral leaves; each peduncle with 3 flowers 
on 2-8 mm long pedicels. Corolla whitish (turning brown on drying), 
1.5—2 mm long, incised up to middle with oblong-ovate subacute lobes. 
Stamens 4. Mericarps glabrous and smooth, about 2 mm long and 
1.5—2.5 mm broad. June to July. 


'This and the following species were described by B.K. Schischkin. 


273 


257 


On stony slopes.— Western Siberia: Irtysh. Endemic. Described 
from Mt. Mramornaya between Nikolaevka and Uspenka. Type in 
Tomsk. 

Note. At present the plant is known only from the one, indicated 
spot. The specimens sent for study from Tomsk, thanks to the courtesy 
of Dr. L.P. Sergievskaya, are well distinguished from the closely related 
species, A. paniculata Bge., widespread in the Altai. 


63. A. tinctoria L. sp. pl. (1753) 104; DC. Prodr. IV, 582; Ldb. 
FI. Ross. II, 398; Boiss. Fl. or III, 44; Schmalh. Fl. II, 10; Kryl. Fl. 
Zap. Sib. X, 2573.—Galium tinctorium Scop. Fl. Carn. ed. II, 1 (1772) 
101.—Ic.: Hegi, Ill. Fl. VI, 1, 206, tab. 114a.—Exs.: Fl. Siles. exs. 
No. 205; Pl. Finl. exs. No. 1350; Fl. Gall. et Germ. exs. No. 657; FI. 
Hung. exs. No. 689. 

Perennial. Rootstock finely funiform, creeping, branched. Stems 
solitary, 30-70 cm tall, straight or ascending, 4-angled, with white 
bands along ribs, glabrous, branched, branches straight, long, upright 
toward tip. Lower leaves in whorls of 6, upper in whorls of 4, unequal, 
linear, (18)30—35(50) mm long, (0.5)1—2(4) mm broad, obtuse, with 
revolute margin, with one vein, glabrous; floral leaves in inflorescence 
paired, very small, at base of cymes elliptic, broad, (1.5)2—3(4) mm 
long, (0.5)1—1.5(2) mm broad, obtuse, glabrous. Inflorescence at end 
of stems and branches, lax on long ternately branched peduncles, 
bearing 3 cymes of 3-6 flowers on each branch. Corolla white, tubular- 
campanulate, 2.5-3 mm long, with tube equaling lobes or 2 times as 
long, tripartite, rarely tetrapartite, lobes elliptic, rounded at tip. Stamens 
3, rarely 4. Anthers longer than filaments, basifixed, barely exposed. 
Styles short, reaching half of corolla tube, with globose stigmas. Fruits 
binate, often one of the mericarps rudimentary, lustrous, wrinkled. 
May to July. 

Meadows, steppe slopes of mountains, forest clearings, open pine 
woodlands.— European USSR: Baltic Region, Upper Volga, Volga- 
Kama, Upper Dnieper, Middle Dnieper, Volga-Don, Volga Region, 
Upper Dniester, Black Sea Region, Lower Don. Western Siberia: Upper 
Tobol. General distribution: Scandinavia, Central Europe, Atlantic 
Europe. Described from Sweden. Type in London. 


64. A. platygalium Maxim. in Bull. Acad. Sc. Petersb. XIX (1874) 
284 (quoad var. alpina Maxim.); Mél. Biol. IX, 267; Palibin in Tr. 
SPb. bot. sada, XVII, 107; Kom. l.c. XXV 484 (only var. alpina 
Maxim.); Kom. and Alis. Opred. rast. Dalnevost. kr. II, 947.—Ice.: 
Kom. and Alis. l|.c. tabl. 283. 


274 


258 


Perennial. Rootstock slender, creeping, woody, branched, red- 
brown. Stem straight, 20-35 cm tall, glabrous, 4-angled, somewhat 
branched upward. Leaves in whorls of 4, lower leaves orbicular-ovate, 
rest ovate, ovate-oblong or elliptic, (14)20—25(28) mm long, (7)10—- 
11(15) mm broad, obtuse, abruptly tapered at base into short petiole, 
thick, with 5 veins, scabrous along margin with isolated, short, upward 
directed setae; elsewhere glabrous, at base with ring of short dense 
hairs; leaves at base of inflorescence smaller; floral leaves in branches 
of inflorescence paired, minute, (1)1.5—2(3) mm long, (0.5)1(2) mm 
broad, resembling cauline leaves in shape and pubescence. Inflorescence 
spicate-paniculate, more or less dense, compact. Peduncles doubly 
ternately branched, together with pedicels glabrous, elongated. Corolla 
white (?), 4-5 mm across, campanulate, with short tube tapered at 
base; lobes 4, oblong, somewhat longer than tube, obtuse or subobtuse. 
Stamens pendent, with filaments longer than anthers, exserted from 
corolla. Style bifid from middle, slightly exserted from flower. Fruits 
binate, subglobose, 3 mm long, 4 mm broad, or with one mericarp, 
glabrous, micro-granulate. Flowering June to July. Fruiting August to 
September. 

Oak groves and mixed forests on mountain slopes.—Soviet Far 
East: Zeya-Bureya (very rarely), Ussuri. Endemic. Described from South 
Ussuri Region (Vaifudin River). Type in Leningrad. 


65. A. maximowiczii Kom. in Tr. SPb. bot. sada, XXXIX (1923) 
109; Kom. and Alis. Opred. rast. Dalnevost. kr. II, 947.—A. 
platygalium var. pratensis Maxim. in Bull. Acad. Sc. Pétersb. XIX 
(1874) 284.—Ic.: Kom. and Allis. l.c. tabl. 284. 

Perennial. Rootstock slender, creeping, reddish. Stem straight, 30— 
100 em tall, glabrous, 4-angled, branched. Leaves in whorls of 4-6, 
oblong-elliptic or elliptic, (23)35—40(53) mm long, (7)9-10(18) mm 
broad, obtuse, gradually tapered at base into short petiole, thin, with 
3-5 veins, with short, soft, erect hairs along margin and beneath along 
veins, with ring of short dense hairs at base; leaves on axis of 
inflorescence in whorls of 4, smaller, in branches of cymes paired, 
much smaller, (2)3(5) mm long, 0.5-1 mm broad. Inflorescence 
paniculate, spreading, very lax, with long axillary branches. Peduncles 
ternately, or doubly ternately, or furcately branched, together with 
pedicels glabrous. Corolla white, pale pink, toward end of anthesis, 3 
mm across, campanulate, with short, broad tube; lobes 4, erect, oblong, 
obtuse, equaling tube or somewhat shorter. Stamens with short filaments 
inserted toward middle of tube. Style short, bifid at tip. Stigmas 
globose. Fruits binate, subglobose, 1.5 mm long, 2 mm broad, rarely 
with one mericarp, glabrous, granular. June to August. 


275 


259 


In oak grooves on mountain slopes and in scrub.—Soviet Far 
East: Ussuri. General distribution: Northeastern China, Korea. 
Described from Posjet Bay. Type in Leningrad. 


Series 3. Aparineae Pobed. Stems clinging. Leaves in whorls of 6— 
8, lanceolate, oblong-lanceolate or oblong-elliptic. Lobes of corolla 
with short, thick incurved tips. 


66. A. rivalis Sibth. et Sm. Pr. Fl. Gr. I (1806) 87.—A. aparine 
auct. non M.B.:: Bess. Fl. Galic. I (1809) 114; Ldb. Fl. Ross. II, 401, 
p-p. Boiss. FI. or. III, 43, p.p.; Schmalh. FI. II, 8, p.p.—Ic.: Syreistsch. 
Fl. Mosk. gub. III, 188; Sibth. et Sm. Lc. II, tab. 117; Rchb. PI. crit. 
Cent. I, tab. XCIII, f. 199.—Exs.: Fl. Boh. and Morav. No. 193 (sub 
A. aparine); GRF, No. 65 (sub A. aparine). 

Perennial. Rootstock slender, creeping, reddish-brown. Stems 
weak, lodged, 60-200 cm tall, banched, 4-angled, nitid, clinging, 
retrorsely spinulose along ribs. Leaves in whorls of 6—8, lanceolate or 
oblong-lanceolate, (17)35(40) mm long, (2.5)5-6(7) mm broad, 
abruptly narrowed at apex into short cusp, gradually tapered at base, 
retrorsely spinulose along margin, sometimes also diffusely spinulose 
on upper surface; floral leaves in inflorescence in whorls of 4 and 
paired, much smaller than those in stems. Bracts not present in all 
flowers; when present bracts often solitary, elliptic or oblong, fine, 
1.5-3 mm long, 0.5—1 mm broad. Inflorescence lax, paniculate, at end 
of stems and upper branches. Peduncles long, several times as long as 
leaves, retrorsely scabrous, ternately branched and bearing 3 simple 
or furcately branched 3-6 flowered cymes. Pedicels equaling flowers 
or shorter, glabrous. Corolla snowwhite, broadly campanulate, with 
short tube; lobes 4, elliptic, broader above with short, thick incurved 
tip. Stamens exserted, with long filaments, inserted in upper margin 
of tube between lobes, 2 times as long as nearly globose small anthers. 
Styles short, included in tube, somewhat partite at tip. Fruits 1.5—-2 
mm long, 1.75—2 mm broad, glabrous, binate, often with one mericarp, 
macrogranulate. Flowering June to July. Fruiting August. 

Banks of rivers and streams, in bottomland deciduous forests, 
groves, among scrub.— European USSR: Baltic Region, Ladoga-Ilmen, 
Upper Volga, Volga-Kama, Upper Dnieper, Middle Dnieper, Volga- 
Don, Volga Region, Black Sea Region, Lower Don. Western Siberia: 
Upper Tobol. General distribution: Central Europe, Balkans—Asia 
Minor. Described from Greece. Type in Oxford. 


67. A. aparine M.B. FI. taur.-cauc. I (1808) 102 and suppl. 105; 
DC. Prodr. IV, 585; Ldb. FI. alt. I, 139 (excl. cit. Besser); Ldb. FI. 


260 


Ross. II, 401, p.p.; auct. non Sibth. and Sm.: Boiss. Fl. or. III, 43, p.p.; 
Schmalh. FI. II, 8; Kryl. Fl. Zap. Sib. X, 2573.—A. rivalis Grossh. FI. 
Kavk. IV, 25.—Galium uliginosum Pall. Ind. taur. (1801) 102.—A. 
rivalis var. schelkownikowiana E. Bordz. in Vish. Kiivsk. bot. sadu, 
XII-XIII (1931) 140.—Iec.: Rchb. Pl. crit. Cent I, tab. XCIII, f. 198. 

Perennial. Stems weak, trailing, clinging, 60-150 cm tall, simple 
or branched, 4-angled, with more or less short, upward directed aciculi. 
Leaves in whorls of 6-8, oblong-lanceolate or lanceolate, (20)30(35) 
mm long, (4)6-7(8) mm broad, abruptly or gradually narrowed at 
apex into short cusp, gradually tapered toward base, retrorsely spinulose 
along margin, sometimes also diffusely spinulose on upper surface; 
floral leaves in inflorescence in whorls of 4—6 and paired much smaller 
than cauline leaves. Bracts, not present in all flowers, when present 
solitary, oblong-elliptic, 2 mm long, 0.5 mm broad, with more or less 
long cusp at tip. Inflorescence lax, paniculate, at ends of stems and 
upper branches. Peduncles long, several times as long as leaves, 
retrorsely scabrous, twice ternately branched, on end branches bearing 
3 simple or furcately branched 3-6 flowered cymes. Pedicels very 
short, equaling ovary or fruit, therefore, more compact than in previous 
species. Corolla snowwhite, tubular-campanulate, with narrow, short 
tube, 2 times as long as lobes; lobes 4, elliptic with short thick incurved 
tip. Stamens on short filaments, 2 times as short as oblong anthers. 
Style short, reaching half of corolla tube, bifid from middle. Fruits 1.5 
mm long, 2 mm broad, glabrous, binate; often with 1 mericarp, 
macrogranulate. June to August. 

In wet meadows, fields, in scrub.— European USSR: Upper Volga 
(adventive in Moscow), Volga-Don, Black Sea Region, Crimea, Lower 
Don. Caucasus: Ciscaucasia, Dagestan, eastern and western 
Transcaucasia. Soviet Central Asia: Tien Shan, Pamiro-Alai. General 
distribution: Iran- Afghanistan (northern part). Described from Caucasus. 
Lectotype in Leningrad. 


68. A. karataviensis Pavl. in Vestn. Akad. nauk. KazSSR, No. 4 
(73) (1951) 95.—A. aparine auct. non M.B.: O. and B. Fedch. Perech. 
rast. Turkest. III (1909) 136; Rgl. and Herd. in Bull. Soc. Nat. Mosc. 
XegNO.gidits 

Perennial. Rootstock slender, creeping. Stems more or less thick, 
strong, erect, 60-120 cm tall, simple or branched, more or less long, 
branches oblique or almost horizontal, 4-angled, lustrous, clinging 
due to retrorse spinules. Leaves in whorls of 6—8, oblanceolate, 
lanceolate or oblong-elliptic, (20)35—40(50) mm long, (3)5—6(8) mm 
broad, at tip abruptly or gradually narrowed into short cusp, gradually 
tapered toward base, retrorsely spinulose along margin, sometimes 


261 


also diffusely spinulose on upper surface; floral leaves in inflorescence 
in whorls of 4—6 and paired, much smaller than cauline leaves. Bracts 
not present in all flowers, when present solitary, oblong-elliptic, 1.5 
mm long, 0.25 mm broad. Inflorescence paniculate, with long, almost 
horizontal lateral branches, lax, cymes few-flowered. Peduncles and 
pedicels retrorsely scabrous, the latter very short, half the length of 
flowers, thick, thickened at top. Corolla bluish, or pale-violet (darkening 
when dry), tubular-campanulate, with more or less broad, short tube, 
equaling lobes or somewhat longer; lobes 4, elliptic with thick short 
incurved tip. Stamens on long filaments, 2—3 times as long as elliptic 
anthers. Style long, equaling tube. Stigma often exserted. Fruit binate, 
often with long mericarp, subglobose, 2 mm long and broad, 
macrogranulate. June to August (Plate XIV, fig. 2). 

Meadow slopes, wet river banks, walnut and juniper forests.— 
Soviet Central Asia: Dzhungaria-Tarbagatai, Syr-Darya, Pamiro-Alai, 
Tien Shan. Endemic. Described from Karatau Range. Type in Alma- 
Ata. 

Note. There are serious errors in the original description of this 
species: the stems of A. karataviensis differ in no way from the stems 
of A. aparine, in which the ribs are also white and sharply prominent; 
the inflorescence is not short, but, on the contrary, considerably longer 
and more spreading than in A. aparine, and the fruit is also finely 
granulate as in A. aparine. 


Section 6. BRACHYANTHAE Pobed. in Addenda XXII, 708.—Sect. 
Cynanchica DC. § Brachyanthae Boiss. FI. or III (1875) 27.—Corolla 
short-campanulate, greenish yellow or yellow. Stigma globose. Leaves, 
especially on branches, fine; branches usually long, almost horizontally 
directed, with large collection of flowers in axils of leaves. 


69. A. humifusa (M.B.) Bess. Pl. Cremen, suppl. III (1814) 4 and 
Cat. pl. Cremen. 17; M.B. FI. taur.-cauc. II, 105; DC. Prodr. IV, 585; 
Ldb. Fl. Ross. II, 401; Boiss. Fl. or. III, 44; Schmalh. FI. II, 9; Grossh. 
Fl. Kavk. IV, 24.—Galium humifusum M.B. l.c. I (1808) 104.— 
Asperula debilis Ldb. Ind. Sem. Horti Dorpat. (1824) 2; DC. l.c. 584.— 

277 A. humifusa y. debilis Schmalh. |.c. 9; Grossh. l.c. 24.—A. humifusa 
a. conferta C. Koch in Linnaea, XVII (1843) 32.—A. conferta (Koch) 
Stank. in Opred. (1949) 579.—A. humifusa B. diffusa C. Koch l.c.— 
A. humifusa var. hirtula Trautv. in Tr. SPb. bot. sada, VII (1880) 
460.—A. humifusa var. trichantha Trautv. in Bull. Soc. Nat. Mosc. 
XXXIX, 2 (1866) 333.—Exs.: Fl. Pol. exs. No. 733; Fl. Cauc. exs. 
No. 218; Pl. exs. Bulg. or. No. 20. 

Perennial. Rootstock slender, creeping, woody, reddish; root thick, 
woody. Stems one-several, prostrate, 20-100 cm long, slender, 


278 


262 


4-angled, more or less densely hairy with long, white, soft hairs, 
especially on lower internodes or scabrous because of setae or mildly 
pubescent, rarely glabrous, lustrous, with numerous, slender, long, 
almost horizontally inclined branches, bearing flowers almost from 
the base of stem, only lowest branches sterile. Leaves in whorls of 6, 
narrowly linear, (5)10—15(23) mm long, (1)2.5-3(5) mm broad, 
cuspidate, with revolute margin and here scabrous because of short 
antrorse setae or hairy on both surfaces with long, white hairs; leaves 
deflexed, appressed to stem, floral leaves (on branches and stems) 
much smaller, (2.5)4—-5(8) mm long, (0.25)0.5(1) mm broad, not 
deflexed, with more compact whorls. Cymes 2-3, rarely 5-flowered in 
axils of all leaves in branches and ends of stems, sometimes gathered 
in corymbose panicle. Peduncles and pedicels short, short-hairy; 
pedicels deflexed in fruit, with one bract at base, not differing from 
floral leaf. Corolla yellow, campanulate, with narrow, short (0.5 mm) 
tube, equaling lobes or shorter, 2-3 mm across; lobes oblong, gradually 
acuminate, deflexed on flowering. Stamens with wide, elliptic anthers 
on filaments equaling or longer than anthers, inserted somewhat below 
the upper margin of tube. Styles bipartite from middle, with unequal 
lobes. Stigmas globose. Fruits minute, 1-1.5 mm long, 1.5—2 mm 
broad, binate, easily splitting into mericarps, broadly elliptic, glabrous, 
very rarely pubescent. May to August. 

On rocks, mountain slopes, calcareous outcrops, plains in mid- 
montane belt, coastal sands and in saline places and as weed.— 
European USSR: Upper Dnieper, Middle Dnieper, Volga-Don, Volga 
Region, Upper Dniester, Bessarabia, Black Sea Region, Crimea, Lower 
Don, Lower Volga. Caucasus: all regions. Soviet Central Asia: Aralo- 
Caspian, Balkhash Region, Dzhungaria-Tarbagatai, Tien Shan (very 
rarely). General distribution: Balkans-Asia Minor, Armenia-Kurdistan, 
Iran. Described from Crimea. Type in Leningrad. 

Note. A highly polymorphic species, within which, as seen from 
the aforementioned synonyms, many varieties and species have been 
described. The pubescence of the stems varies; the leaves, in varying 
degrees, are scabrous or densely pubescent on both surfaces with long, 
white hairs; the corolla is glabrous or pubescent on the outside; the 
fruits are glabrous or setose. 


70. A. insuavis Pobed. sp. n. in Addenda XXII, 708.—A. humifusa 
auct. fl. As. med. non Bess.—A. humifusa 6. aspera Regel and Herder 
in Bull. Soc. Nat. Mosc. XL, 1 (1867) 8. 

Perennial. Rootstock slender, creeping, woody, reddish; root thick, 
woody. Stems one-several, suberect, 10-100 cm long, more or less 
thick, 4-angled, glabrous, lustrous or scabrous because of short retrorse 


279 


263 


setae, rarely pubescent with scattered, white, soft hairs, with numerous 
slender, longer, almost horizontally inclined branches, often repeatedly 
branched, floriferous almost from the base, only lowest branches sterile. 
Leaves in whorls of 6, linear, (10)25—30(40) mm long, (1)3—4(5) mm 
broad, cuspidate, flat or with revolute margin, glabrous or scabrous 
along margin from short antrorse setae; leaves deflexed, appressed to 
stem; floral leaves (on branches and stems) considerably finer, (2.5)5— 
6(8) mm long, (1)1.5(2.5) mm broad, not deflexed, with more compact 
whorls. Cymes 2-3, rarely 5-flowered in axils of all leaves in branches 
and end of stems, sometimes, forming corymbose panicles. Peduncles 
and anthophores, short, glabrous or rarely pubescent; latter deflexed 
in fruit with usually one bract at base, not differing from floral leaves. 
Corolla 2—2.5 mm across, yellow, campanulate, with tube somewhat 
longer than lobes (1 mm long), rarely equaling lobes; lobes oblong, 
obtuse, with small deltoid incurved tip. Anthers elliptic, almost equaling 
filaments, inserted somewhat below upper margin of tube. Style bifid 
from middle, with unequal lobes, Stigma globose. Fruits 1.5 mm long 
and 2 mm broad, binate, easily splitting into elliptic mericarps, 
glabrous, rarely pubescent, microgranulate. June to September. 

On mountain slopes, in rocks in steppe and deserts, rarely in 
weedy places.—Soviet Central Asia: Balkhash Region (very rarely), 
Dzhungaria-Tarbagatai, mountainous Turkmenia, Syr-Darya, Pamiro- 
Alai, Tien Shan. General distribution: Iran. Described from southern 
Tadzhikistan. Type in Leningrad. 


Section 7. Oppositifoliae Schischk. sect. n.— Flowers 4-merous in 
cymes at tip of stem and branches. Corolla dirty pink or whitish, tube 
2-3 times as long as limb. Perennials with woody base. Leaves 
opposite. 


71. A. oppositifolia Rgl. et Schmalh. Opis. nov. vidov v 
Puteshestv. v. Turkestan A.P. Fedchenko. Vyp. 18 (1881) 42.—O. 
and B. Fedchenko, Perech. rast. Turk. III, 136.—A. sordide-rosea M. 
Pop. in E. Korov., M. Kult. and M. Popov. Description of new plant 
species, collected in Turkestan (1916) 73.—A. albiflora M. Pop. (1916) 
l.c.—Exs.: Bornm. Pl. turkest. (1913) No. 531. 

Perennial. Base woody. Stems numerous, ascending or straight, 
branched, indistinctly 4-angled, glabrous or short-pubescent in upper 
part, 5-25 cm tall, Leaves opposite, linear or linear-lanceolate, 
subobtuse or acute, finely scabrous along margin (under a magnifying 
glass), 1-1.5 cm long and 1-3 mm broad. Cymes at end of stems and 
branches. Peduncles often with short isolated pubescence. Bracts 
herbaceous, resembling leaves, equaling glabrous or shortly pubescent 


264 


pedicels in fruit, at anthesis shorter than pedicels. Pedicels 2.5 mm 
long in fruit. Flowers 4-merous. Corolla whitish, clavate-tubular, 
sparsely pubescent (under a magnifying glass) or glabrous, tube 2 or 
almost 3 times as long as deltoid or ovate lobes. Stigma usually globose. 
Mericarps densely setose-hairy, setae almost equaling mericarp in 
diameter. July to August (Plate XV, fig. 1). 

On gypsum sandstones, rocks and stony slopes at altitudes of 
1800-2400 m.— Soviet Central Asia: Pamiro-Alai, Tien Shan (Angren 
Range), Chatkal, etc. Endemic. Described from Pamiro-Alai. Type in 
Leningrad. 

Note. 1. M.G. Popov, I.c. (1916), believes that A. oppositifolia 
Rgl. and Schmalh. represents a species aggregate and that, in accordance 
with the rules of botanical nomenclature, the name A. oppositifolia 
Rgl. et Schmalh. should be abolished. Such an assertion is 
totally erroneous, since in the event that one species is divided into 
two, the original name must be preserved for one of them. 
A. oppositifolia Rgl. et Schmalh. varies quite widely, but a fundamental 
characteristic of this species are the long, persistent hairs on the ovary 
and fruits. With respect to the pubescence of the corolla tube as well 
as the length of the tube and limb, these features are not stable. The: 
color of the corolla also varies from whitish to dirty pink. From the 
foregoing, we consider it possible to preserve the name A. oppositifolia 
Rgl. et Schmalh., understanding it in a somewhat broader sense. 

Note 2. From Kugitang in valley of Tangi Dara, we have one 
incomplete herbarium specimen with young flowers collected and 
named by M.G. Popov as Asperula kelleri M. Pop., renamed 
subsequently by O. Tschernova as A. pauciflora Tschernova. The 
pubescence of the ovaries places it in between A. oppositifolia Rgl. et 
Schmalh. and A. popovii Schischk. In view of the insufficient material, 
we consider it inappropriate to distinguish this plant as a separate 
species. 

Note 3. Among the plants preserved in the Herbarium of the V.L. 
Komarov Botanical Institute of the Academy of Sciences under the 
name A. oppositifolia Rgl. et Schmalh. there are stunted specimens, 
not exceeding 4—6(10) cm in height. These specimens are known 
predominantly from Darvaz in the Vakhsh Range. B.A. Fedtschenko, 
in the herbarium, annotated these plants as a distinct species—A. 
trajectorum B. Fedcz., whereas P.N. Ovchinnikov referred to the species 
as A. fedtschenkoi Ovcz. Further investigation is necessary. 


280 


265 


72. A. popovii Schischk. nom. nov.—A. oppositifolia ssp. sordide- 
rosea, M. Pop. in Preface to Byullet. Sr. Az. gos. univers. Vyp. 14 
(1926) 18.—Exs.: H.F.A.M. No. 250. 

Perennial. Base woody. Stems numerous, ascending at base, 
thereafter straight, branched, with obliquely erect branches, glabrous, 
slender, 4-angled, 20-40 cm tall. Leaves opposite, linear, glabrous, 
with one vein, acute or subobtuse, 0.5—1.5(2) cm long. Cymes at ends 
of stems and erect short branches, usually 3-flowered, of these middle 
ones sessile, lateral ones on short pubescent pedicels. Bracts fine, 
herbaceous, puberulent, resembling leaves, but much smaller. Flowers 
4-merous. Corolla dirtyish pink, about 4.5 mm long, tube usually 
covered with short, isolated white hairs, incised up to 1/3 into ovate, 
obtuse teeth. Ovary covered with very short, sometimes less noticeable, 
isolated, white hairs. Mericarps densely pubescent with straight, long 
hairs. July to August. 

In crevices of rocks up to altitudes of 2400 m.—Soviet Central 
Asia: Tien Shan (Chimgan, Ugam. Talas Ala Tau ranges), Pamiro- 
Alai (Alai Range). Endemic. Described from western Tien Shan. Type 
in Tashkent, cotype in Leningrad. 


73. A. laevis Schischk. sp. n. in Addenda XXII, 709. 

Perennial. Entire plant glabrous, woody at base. Stems numerous, 
ascending or straight, branched, with obliquely erect short branches, 
4-angled, 20-35 cm tall. Leaves opposite, linear, with one vein, acute 
or obtuse, 1-2 cm long and 0.5—1 mm broad. Cymes at ends of stems 
and branches forming narrowly paniculate inflorescence; floral leaves 
herbaceous, lanceolate or narrowly ovate, acute, 2-7 mm long and 
0.5—1 mm broad. Pedicels 1-5 mm long, filiform. Corolla light pinkish, 
about 5 mm long, tube 2 times as long, narrowly ovate, subobtuse 
lobes. Mericarps oblong, glabrous and smooth, about 2 mm long. July 
to August (Plate XV, fig. 2). 

On stony slopes in gypsum limestone, in rocks at altitudes of 
1500-3500 m.— Soviet Central Asia: Pamiro-Alai. Endemic. Described 
from Darvaz. Type in Leningrad. 


Section 8. SHERARDIANA DC. Prodr. IV (1830) 581; Boiss. FI. or. 
III, 30; Schumann in Pflanzenfam. IV, Abt. 4-5, 149.— Inflorescence 
capitate. Corolla 4-merous, blue, dark blue, rarely white, lobes without 
appendage. Annual herbs. 


Series 1. Arvenses Schischk.—Corolla tube up to 7 mm long, 
lobes of corolla obtuse or subobtuse. Bracts usually longer than flowers. 


— 


266 


Economic importance. In the studies of D.I. Mugalinskaya (Izv. 
Azerb. fil. Akad. nauk SSSR (1943) No. 2, 34-39), some members of 
the series Arvenses Schischk. were found to contain a milk-curdling 
enzyme. Richest in the enzyme are the leaves, inflorescence and 
especially the fruit; powders of these, which contain the enzyme, are 
used for preparing cheese. The author of this work identifies the species 
of woodruff [Asperula] used as A. arvensis L. According to our studies, 
in Azerbaidzhan Asperula setosa Jaub. et Spach (and not A. arvensis 
L.) is widespread and, in all probability, Muganlinskaya had worked 
with this species. Evidently, all members of the series Arvenses 
Schischk. contain the same enzyme. 


74. A. arvensis L. Sp. pl. (1753) 103; Ldb. FI. Ross. II, 397 ex 
p.; Boiss. Fl. or. III, 30; Schmalh. Fl. II, 7, part.; Grossh. Fl. Kavk. 
IV, 23, ex p.—A. arvensis var. vulgaris Trautv. in Tr. SPb. bot. sada, 
VII, 2 (1880) 459.—Ic.: Rchb. Ic. Fl. Germ. XVII, tab. 1177. 

Annual. Stem simple or branched at base, with obliquely upward 
directed branches, 4-angled, glabrous, usually scabrous along edges, 
often whitish, 20-40 cm tall. Lower leaves persistent (cotyledonary 
leaves perishing early), opposite, broadly ovate, abruptly tapered into 
short, broad petiole, 10-12 mm long and 6-7 mm broad, tip rounded, 
often violet colored; remaining leaves whorled, in whorls of 6-8, 
lanceolate or lanceolate-linear, tapered at base, sessile, obtuse or 
subacute at apex, with one middle, prominent, often slightly scabrous 
vein beneath, margin somewhat revolute and with short, acute, whitish 
aciculi, leaves of 2 upper whorls usually with long, rigid, white setae. 
Flowers in capitate inflorescence at end of stem and branches, 
surrounded by involucre of numerous, lanceolate-linear, scabrous leaves 
7-10 mm long and 1-1.5 mm broad, with obliquely standing, 1—1.5 
mm long white setae along margin, especially in lower half; each 
flower having ciliate bracts somewhat shorter than leaves on involucre. 
Corolla light blue or, rarely white, glabrous, about 6 mm long, with 
short lobes. Mericarps globose, about 2.5 mm in diameter, finely 
tuberculate (under a magnifying glass). May to June. 

In fields, around dwellings, in forest clearings, parks — European 
USSR: Middle Dnieper (vicinity of Kiev, adventive), Volga-Don 
(vicinity of Kharkov, adventive), Lower Don (Novocherkassk, 
adventive), Upper Dniester (?), Crimea. Caucasus: Ciscaucasia, eastern 
Transcaucasia (very rarely adventive). General distribution: Central 
and Southern Europe, Balkans-Asia Minor, Armenia-Kurdistan, Iran(?), 
North Africa(?). Described from France and Germany. Type in 
London. 


282 


267 


Note. Study of the herbarium material preserved in the Herbarium 
of the Botanical Institute, Academy of Sciences, USSR did not provide 
a clear picture of the distribution of this species within the USSR. 
From the Crimea and Ciscaucasia, there are numerous specimens of A. 
arvensis L., whereas from the more northern regions of European USSR 
there are only a few specimens (e.g., from Kharkov, Novocherkassk). 
Schmalhausen (1.c.) reports A. arvensis L. for Kiev. It must be observed 
that all the aforementioned localities are based on the collections of 
early collectors. Collections of this species from the reported or any 
other localities in the European part of the USSR have not been added 
to the Herbarium of the Botanical Institute, Academy of Sciences, 
USSR, in recent times. From eastern Ciscaucasia, also, there are only 
meager collections of A. arvensis L. from Shamkhor, Belyi Klyuch, 
and Zarzma Station, evidently collected in tilled fields or around 
railway stations. In Transcaucasia, another species, A. setosa Jaub. et 
Sp., is widespread. 


75. A. setosa Jaub. et Sp. Ill. Pl. or (1842-1843) 152; Boiss. Fl. 
or. III, 30; Nevskii in Tr. Bot. inst. ser. 1, [V (1937) 275.—A. arvensis 
var. ambigua Trautv. in Tr. SPb. bot. sada, VII, 2 (1880) 460.—A. 
arvensis Grossh. Fl. Kavk. IV, 23, ex. p.—Ie.: Jaub. and Sp. l.c. tab. 
82. 

Annual. Stem solitary, rarely stems numerous, whitish, 4-angled, 
more or less scabrous along edges, simple or sometimes branched 
from base, 8—30 cm tall. Lowermost leaves opposite, on short petioles, 
broadly ovate, 6-7 mm long and 4—5 mm broad, falling early, glabrous, 
upper leaves whorled, 4-8 leaves in whorl, sessile, lanceolate-linear, 
obtuse, tapered toward base, with one prominent vein beneath, usually 
with short hairs along margin and vein, 1-2 cm long and 1-2 mm 
broad. Flowers in capitate inflorescence at end of stem and branches, 
inflorescence 1—1.5 cm in diameter, surrounded by involucre leaves 
and bracts. Involucre leaves similar to cauline leaves, but smaller, like 
bracts, with long usually antrorse, white or lightly violet colored hairs 
along margin, 2-3 mm long. Corolla cylindrical, light blue or white, 
glabrous, about 6 mm long, equaling, somewhat shorter or longer than 
involucre leaves. Mericarps globose, 1-1.2 mm in diameter, with short, 
often numerous, white hairs. Flowering May to June. Fruiting July. 

Grassy and stony slopes, shingles, gardens, vineyards, and fields 
up to an altitude of 2400 m.— Caucasus: Dagestan, eastern, southern 
and western (very rarely) Transcaucasia, Talysh. Soviet Central Asia: 
Tien Shan (western part), Pamiro-Alai, mountainous Turkmenia. 
General distribution: Armenia-Kurdistan, Iran, Afghanistan, Asia 
Minor. Described from Iran (Mt. Elvend). Type in Paris. 


283 


268 


Note. We refer Caucasian and Central Asian plants, usually 
identified as A. arvensis L., to A. setosa Jaub. et Sp., although the fruit 
of our plant differs sharply in having a smaller size, the absence of 
tubercles and the presence of tiny, linear, white bands on the fruit, 
depicted by the authors of this species in the plate. Ignoring the drawing 
of the fruit by Jaubert and Spach, Boissier (Boiss. Fl. or. III, 30), in 
describing the fruits of A. setosa Jaub. et Sp., indicates that the presence 
of white, linear bands, is extremely characteristic also of the Central 
Asian and Caucasian plants. 


Series 2. Azureae Schischk.—Corolla tube 8-14 mm long, 3-5 
times as long as limb; corolla lobes subacute. Bracts considerabiy 
shorter than flowers. 


76. A. azurea Jaub. et Sp. Ill. pl. or. (1842-43) 151; Walp. Repert.. 
VI (1846-1847) 24; E. Regel, Ind. sem. Horti Petrop. 52; G. Woron. 
and A. Schelkown. Herb. Fl. Cauc. Fasc. V, 21.—A. azurea setosa 
Rgl. in Gartenfl. XV (1866) 323, non A. setosa Jaub. et Sp.; in Vestn. 
Ross. obshch. sadov. in SPb. (1867) 45.—A. orientalis Boiss. et Hoh. 
in Boiss. Diagn. ser. 1, 3 (1843) 30; Boiss. Fl. or. HI, 31; Grossh. FI. 
Kavk. IV, 23.—A. orientalis var. longiflora Albov. in Tr. Tifl. bot. 
sada, I (1895) 121.—A. arvensis var. orientalis Trautv. in Tr. SPb. 
bot. sada, VII, 2 (1880) 460.—Ic.: Jaub. and Sp. l.c. tab. 82, A; Rgl. 
l.c. (1866), tab. 523, f. 4; E’. Regel. l.c. (1867) tabl. 4.—Exs.: Herb. 
Fl. Cauc. No. 243. 

Annual. Stems solitary or several, 10—20 cm tall, 4-angled and 
along margins in lower part often with very short, acicular hairs, simple 
or branched, often almost from base. Lower leaves falling early, 
opposite, broadly ovate or almost orbicular, 4-7 mm long and 2-5 
mm broad, on short petioles, glabrous, ends obtuse or rounded; upper 
leaves whorled, 4 or 6 leaves in whor!, lanceolate or linear-lanceolate, 
tapered at base, sessile, subobtuse or acute, with one prominent vein 
beneath, sometimes scabrous along margin and middle vein, 1.5—3 cm 
long and 2-5 mm broad. Leaves of upper whorls with erect, long, 
white setae. Flowers in capitate inflorescence at ends of stem and 
branches, surrounded by numerous linear, subacute bracts, usually 
scabrous with short hairs and, moreover long, white, erect setae, 
predominantly in lower half along margin. Corolla cylindrical, 8—14 
mm long, light or dark blue, very short velutinous, with 4 short, ovate- 
deltoid lobes. Mericarps smooth, covered with numerous, short, whitish 
bands. Flowering April to June. Fruiting July. 

Stony slopes, pine forests up to an altitude of 1000 m.— Caucasus: 
Ciscaucasia, eastern and western Transcaucasia. General distribution: 


284 


269 


Asia Minor, Armenia-Kurdistan, Iran. Described from Asia Minor. 
Type in Paris. 

Note. This plant was described in the same year (1843) as a new 
species under two names: A. azurea Jaub. et Sp. and A. orientalis 
Boiss. et Hoh. According to the international rules of botanical 
nomenclature, in the absence of an indication of a more exact 
publication date of the respective books, priority will be given to the 
name which was accepted by the earliest subsequent authors. In the 
present case the name A. azurea was used by Walpers (Walp. l.c.) and 
E. Regel (Rgl. l.c.), whereas A. orientalis came into use only after the 
publication of the third volume of “Flora Orientalis” by Boissier (1875). 

Economic importance. In bloom, species is one of the most 
beautiful annuals, easily cultivated and, as early as 1866, was 
recommended by E. Regel, for wide propagation as an ornamental 
plant for open grounds. 


SPECIES OF INDISTINCT AFFINITY 


77. A. taurica Paczoski in Zap. Novoross. Obshch. estestv. XV, 
1 (1890) 76; Lipsky in Tr. SPb. bot sada, XIII, 300. Schmalh. FI. II, 
8.—A. cretacea Ldb. FI. Ross. II (1844-1846) 399, non Roem, et 
Schult.—A. hexaphylla auct. non All.: Georgi, Beschr. Russ. Reich. 
III, 4 (1800) 728; Fedtsch. et Fler. Fl. Evrop. Ross. 906.—Exs.: GRF, 
No. 266; Herb. norm. (Dorfler), No. 5086. 

Perennial. Rootstock slender, branched, brown. Stems numerous, 
branched or simple, 6-20 cm tall, slender, 4-angled, glabrous. Leaves 
in whorls of 6, lowermost leaves in fours, narrowly linear, (10)17— 
18(24) mm long, (0.5)1(15) mm broad, mucronate, sessile with one 
vein, thicker; floral leaves at base of inflorescence somewhat smaller 
than cauline leaves. Bracts solitary, considerably smaller than leaves. 
Cymes terminal in threes on stems and branches, ‘simple, two-five- 
flowered, peduncles and pedicels glabrous. Corolla white, 4 mm across 
with short tube, equaling lobes or somewhat longer, with 4 oblong 
obtuse lobes, slightly tapering toward tip. Filaments somewhat longer 
than anthers, inserted below upper margin of tube; anthers included in 
tube. Styles short, 2/3 connate; stigma globose. Fruit often with one, 
almost globose mericarp, 2.5 mm long and 2 mm broad or binate, 
glabrous, granular on surface. May to July. 

On limestone and chalky mountain slopes, stony slopes.— 
European USSR: Crimea. Caucasus: Ciscaucasia. Endemic(?). 
Described from Crimea. 


285 


270 


Note. In the classification of the genus, this species stands apart. 
The species A. hexaphylla All., described from Italy, to which our 
Crimean species has often been referred, is close to it; however, the 
Italian species is well distinguished from the Crimean in having long 
styles exserted from the corolla and a tube two times longer than its 
lobes. Which species Kozo-Polyansky [in Karsten, Veg. Bild. XXII 
(1931) t. 10] had in mind under the name A. cretacea Schlecht. is still 
unknown for lack of material of this species, and also because of the 
absence of any diagnosis either in Schlechtendahl, who only provided 
the name on an herbarium label, or in Kozo-Polyansky, who was mainly. 
interested in the occurrence of this plant in the chalks along the Tikhaya 
Sosna River. The photograph in Kozo-Polyansky’s paper does not 
allow one to distinguish the species. 


78. A. gracilis C.A.M. Verzeichn. Pfl. Cauc. (1831) 52; Ldb. FI. 
Ross. II, 397; Boiss. Fl. or. III, 43: Grossh. Fl. Kavk. IV, 24. 

Perennial. Stem slender, filiform, 5-15 cm long, branched in upper 
half, terete or 4-angled, with poorly expressed ribs. Leaves in whorls 
of 4, elliptic, 5-8 mm long, 2.5-3 mm broad, obtuse, on very short 
pedicels, glabrous, barely scabrous along margin, uppermost leaves 
paired, finer. Cymes, 3-4 flowered, simple, terminal. Peduncles short, 
glabrous. Pedicels several times as short as flowers. Corolla white, 
infundibuliform, almost glabrous, 2-3 mm across, with 4—5 elliptic, 
obtuse lobes. Anthers narrow, long, 2 times as long as filaments. Style 
bifid from middle half, shorter than tube, included. Fruit glabrous, 
smooth. July. 

In wet areas in alpine zone.— Caucasus. Described from Caucasus 
(Malka River). Type in Leningrad. 


GENUS 1390. Leptunis'* Stev. 


Stev. in Bull. Soc. Nat. Mosc. XXIX (1856) 366. 

Flowers bisexual, terminal, on long trichoid pedicels; lateral 
flowers sessile, with bracts at base. Calyx limb absent. Corolla tubular, 
pale-carneous, lobes of limb oblong, obtuse, one-third the length of 
tube, puberulent outside. Stamens 4, anthers lanceolate, black. Ovary 
bilocular; styles equaling tube, bifurcate at tip, with globose stigmas. 
Mericarps clavate, incurved at tip, immature mericarps puberulent, 
subsequently glabrous. Annuals, with filiform, whorled leaves, 10-16 
in a whorl. 


‘Treatment by B.K. Schischkin. 
*From the Greek Jeptos, meaning thin. 


286 


271 


In this genus there are, apparently, 2-3 species. The number of 
species is still not clear because of the absence of ripe fruits on the 
available herbarium specimens of closely related species found in Iran. 


1. L. trichoides (J. Gay) Schischk. comb. n.—L. tenuis Stev. l.c. 
(1856).—Asperula trichoides J. Gay. in DC. Prodr. IV (1830) 582; 
Boiss. Fl. or. IfI, 31; B. Fedtsch. Rast. Turkest. 710; Grossh. Fl. Kavk. 
TN 23. 

Annual. Stem 8-25 cm tall, terete, filiform, glabrous, erect, 
trichotomously branched usually from middle or above, with trichoid 
branches. Lowermost leaves linear, opposite, early falling; cauline 
leaves filiform-subulate, acute, stiff, slightly scabrous, 10-16 in a 
whorl, 4 or 2 in a whorl on branches; uppermost leaves solitary, 2— 
3.5 cm long. Pedicels patent above, trichoid, linear, lateral, if present, 
very short (flowers almost sessile). Corolla pale-carneous, under 
magnification appressed puberulent, 3 mm long, with short, cylindrical 
tube, distinctly expanded in throat, with limb of oblong, obtuse lobes, 
half the length of tube. Anthers ovate, black. Styles equaling, bifurcate 
at tip. with globose, stigmas. Mericarps obovate, sparsely appressed 
puberulent, 1.5 mm long and 1 mm broad, with thin, whitish, linear, 
bands (Plate XV, fig. 4). 

On stony rubbly, clayey and sand slopes and shingles.— Caucasus: 
Southern Transcaucasia, Talysh. Soviet Central Asia: Dzhungaria- 
Tarbagatai (only Dzhungarian Ala Tau), Tien Shan (western part), 
Pamiro-Alai, mountainous Turkmenia. General distribution: Iran. 
Described from specimens grown from seeds sent by Fisher from 
northern Iran. Type in Geneva. 


GENUS 1391. Microphysa’” Schrenk 


Schrenk in Bull. Phys.-math. Acad. Sc. Pétersb. II (1844) 115. 

Flowers bisexual, 4-merous. Calyx without lobes. Corolla almost 
infundibuliform, with short, narrow tube. Filaments inserted somewhat 
below upper margin of corolla tube. Ovary bilocular, each locule one- 
seeded. Fruit dry binate. Mericarps included in vesicular calyx: latter 
connate at top with coarsely granulate surface. Inflorescence paniculate. 
Perennial herb with branched, strong erect stems. 

A monotypic genus, in habit similar to species of the genus Galium, 
in the form of the flower not differing from the genus Asperula, in the 


'Treatment by E.G. Pobedimov. 
7From Greek micros—meaning small, and physa—a bubble. 


287 


272 


fruits, with two mericarps enclosed in a common covering, having a 
resemblance to the genus Rubia. 
Type of genus: Microphysa elongata (Schrenk) Pobed. 


1. M. elongata (Schrenk) Pobed. comb. n.—M. galioides Schrenk 
in Bull. phys.-math Acad. Sc. Pétersb. II (1844) 116.—Asperula 
elongata Schrenk, Enum. pl. nov. (1841) 58; O. and B. Fedtsch. Perech. 
rast. Turkest. III, 136. 

Perennial. Root creeping, reddish. Stem erect, branched almost 
from base, with straight branches, erect above or simple, 30-70 cm 
tall, scabrous along ribs due to slanting spinules. Leaves in whorls of 
4, upper ones on branches often opposite, linear-lanceolate, (33)40- 
50(60) mm long, (1.5)4—5(12) mm broad, acute or obtuse, with one 
vein, scabrous along margin in upper part and beneath along vein due 
to spreading spinules, almost sessile or with very short petioles; floral 
leaves opposite, (5)7(12) mm long, (0.5)1.5(3) mm broad. Inflorescence 
terminal, paniculate, corymbose, more or less dense; cymes many- 
flowered. Peduncles doubly furcately or ternately branched, of the 
first order more or less elongated, remaining short pedicels, covered 
with scattered spinules. Corolla white, 3.5-4 mm across, with. 
infundibuliform tube and with oblong, shortly acuminate lobes. 
Filaments equaling almost half lobes of corolla; anthers large. Style 
bipartite in upper 1/3; stigma globose. Fruit binate, included in calyx; 
fruiting calyx vesicular, green, surface granulate, 2.5 mm long and 
broad, connate at top and adnate only to back of fruit, rest free, 
mericarps very small, 1 mm long and broad, orbicular, compressed, 
flat. June to July (Plate XXI, fig. 4). 

In meadows along banks of rivers and lakes, in foothills and 
valleys.—Soviet Central Asia: Aralo-Caspian. Balkhash Region, 
Dzhungaria-Tarbagatai, Tien Shan. Endemic. Described from foothills 
of Labassa (Dzhungarian Ala Tau). Type in Leningrad. 


GENUS 1392. Galium!? L. 


Flowers bisexual, rarely unisexual, monoecious, with lobes of calyx, 
4-merous, rarely 3-merous. Corolla rotate, lobes of corolla connate at 
base or rarely up to 1/3 the length. Stamens with filaments inserted at 
top, very rarely to lower margins of connate part of corolla, in between 


1Treatment by E.G. Pobedimova. 
*From the Greek gala, meaning milk, because these plants promote the coagulation 
of milk. 


288 


213 


lobes. Styles 2, free or often more or less connate. Fruits binate, 
coriaceous, splitting on ripening, or mericarp single on account of 
partial or complete lack of development of the second, glabrous or 
pubescent, smooth, with granulated or rugose surface, sometimes 
swollen. Inflorescence—terminal panicles or cymes on stems and 
branches or axillary, sometimes reduced to solitary flowers. Leaves in 
whorls of 4-14, uppermost floral leaves usually paired or solitary, 
sometimes having bracts. Perennial or annual herbs with simple or 
branched, strong or weak, lodged stems. 

The genus is extremely widespread over the globe and includes 
about 400 species. 

This genus has not been treated monographically to date. A 
systematic treatment is difficult for the following reasons: in the genus 
Galium, often species belonging to different sections are similar in 
habit; at the same time many of its widely distributed species are 
highly polymorphic, but often youthful and weakly differentiated 
geographically. Polymorphic species (e.g., G. boreale L. and G. 
septentrionale Roem. et Schult.) have to be studied with mass 
collections using the cytological method of investigation in order to 
establish their taxonomic significance. On the other hand, ancient 
species of this genus, with disjunct ranges, appear to be extremely 
stable morphologically and do not produce young races where one 
would expect them (e.g., G. paradoxum Michx. in the Urals and the 
Far East). In the genus there often is hybridization between both closely 
related species and species from different sections. A systematic 
treatment of the genus has not been worked out; it differs more or less 
in all the floras. We shall not attempt here to provide a new systematic 
treatment, because we consider this to be possible only after a 
monographic treatment of the genus, but it becomes necessary to 
introduce certain changes to the circumscriptions of sections as well as 
to add new sections for species clearly lying beyond the earlier 
established groupings. 

Economic importance. Certain species of this genus are known 
to produce pigments; e.g., G. verum L. is used to obtain red dye. The 
roots contain dyestuff of the anthroquinone series, chiefly flavopurpurin 
and anthropurpurin (Pogozhev in the journal Za. rekonstruktsiyu 
tekstilnoi promyshlennosti (1934) No. 5, 35). In homeopathy, an extract 
from the fresh flowering herbs is used (G. aparine L., G. mollugo L. 
and G. verum L.) In Tibetan medicine, the roots of G. aparine L., G. 
mollugo L. and G. boreale L. are used. [Stankov and Kovalevskii, 
Nashi lekarstvennye rasteniya (Our medicinal plants) (1952) 110]. The 
herb G. verum L. contains 0.21% of a crystalline glucoside; the herb 
G. triflorum Michx. contains coumarin and hisperidin [Wehmer, Die 


289 


274 


Pflanzenstoffe, 2 (1931) 1181-1182]. G. verum is a nectariferous plant: 
it yields a moderate quantity of nectar [Slovar’-spravochnik pchelovoda 
(Manual for Apiculturists) (1955) 269]. 


1. 


+ 
D 


D4 


00 4 


Leaves in whorls of 4, rarely middle cauline in 4—6, remaining in 


DA) ecatged ses OW rh Dian Ve, Ee LR AY, Seenees 2 
Reaves ii whorlsioh GUO ye Pe Lea eat itttocdasstovct vncreactenaees $). 
Middle cauline leaves in whorls of 4 or 6, remaining in 2 or 4 ... 
PCI. SR, ARE Pe NE DNB NOE a Roane. amet rates canoe ta sente 3. 
All iexiebci itn WHOTISMOF Sie PISA EM. Leese lick. See ceece sete eokasntage 9. 


Flowers solitary, almost sessile in axils of leaves. Mericarps narrow, 
oblong, their length exceeding breadth. Small annual herbs 
[Section Asperae (Moench) DC.] ..........cccccecccesseeeseeeenseeeeneeeees 4. 
Flowers in branched, few-flowered cymes. Mericarps spherical. 
Marge’ perenmialplarttsie 20-222 0s5.. ksvedss.. deters corer tees acetone sere eens 6. 
Pedicels shorter than or equaling fruit, straight, erect ................. : 
Pedicels somewhat longer than fruit, recurved so that mericarps 
nearly pressed against stem. Stem glabrous ...............::eeeeee 
fine RE ee WO Dey Ne 15. G. apsheronicum Pobed. 
Leaves reflexed in fruit. Flowers almost sessile; pedicels thick, not 
longerthan firmitqc! Sagi ke ae. 14. G. verticillatum Danth. 
Leaves not reflexed in fruit. Flowers on slender, filiform, simple or 
furcate, slightly arcuate, horizontal pedicels 2—2% times as long as 


EU OES eel eae 2 16. G. nupercreatum M. Pop. 
Fruit and ovary with long, white, shining, hooked hairs. Leaves 
AILS TOMAS WAI EE ce. «Benes eee, ER asst, Send Seeeerenae Senaencersseecceeeas die 
Fruit and ovary granular-tuberculate. Leaves obtuse, without mucro 
to Rad) ARR, SE RE ESOC SIR. et tete cease. Adeenes res 8. 
Cymes terminal, in axils of upper leaf whorls. Leaves oblong-obovate 
BEALL Rte Te Shane AEE Sees 3. G. trifloriforme Kom. 


Cymes axillary. Leaves oblong-lanceolate ... 2. G. triflorum Michx. 
Leaves oblong-lanceolate, (12)20(25) mm long, (1.5)3(5) mm broad, 
scabrous along margin due to retrorse spinules. Fruit midlly granular- 
tuberculate, often rugose. Stem thick, swollen at nodes ... 
segencpvegh? ox tieeek.. SAOURe, SOT OE IIS, OT, 37. G. elongatum Presl 
Leaves linear, (7)20(30) mm long, (1)2(3) mm broad, scabrous 
along margin due to antrorse spinules. Fruits densely granular- 
tuberculate. Stem slender, not swollen at nodes ............:.ssceseeeeeee 
RRA, DE AR, Se ee Ne, Sees 38. G. krymense Pobed. 
Cymesvanillary estecc62 5.2220. 220 ROR Ea, 10. 
Cymes terminal and in axils of upper leaves only or forming 
paniculate, inflorescemee sx. 20. :ohsstss-cene Rete te aeeee eee eteee 23" 


290 


10. 


11. 


12) 


13) 


16. 


17(11). 


19: 


275 


Cymes single-flowered, with peduncles longer than leaves, erect 
in fruit. Flowers bisexual. Leaves in whorls of 4, 2 smaller than 
Othertwoitwee es ek ROR, 1. G. soongoricum Schrenk. 
Cymes 2-6 flowered, with short peduncles, shorter than leaves, 
recurved in fruit, hidden between leaves. Flowers polygamous, 
unilocular. Fruit with one, more or less globose, mericarp. (Section 


CriciataDG RESO KOR. RE BO SE, 11. 
Gymieshwithout bracts 22. Eo ae Pe... BE 12. 
COUSIN) Cele ts eee ee A neers Sc Sen 17. 


Leaf whorls closely-packed all over stem. Leaves broadly cordate, 
reflexed after anthesis. Fruits covered with minute papillary hairs 
PR SEAN LEY, Sate yh ty eRe ee crt: 28. G. cordatum Roem. and Schult. 
Leaf whorls more or less scattered. Leaves elliptic. Fruits glabrous 
or covered with scattered, long hairs ..........cccescsesseeseeeeeeeeeee 13. 
Leaves (2)5(11) mm long, (0.75)2(5) mm broad. Stem retrorsely 
spinulose. Whorls of leaves widely scattered ..............::ccseseeeees 
pe ieein Rho oetee Hae etenrerdd teye 29. G. pedemontanum (Bell!.) All. 
Leaves 6-25 mm long, 2.5—9 mm broad. Stem glabrous or patently 
pubescent. Whorls of leaves CIOSEF ...........:ccscccseeesesteesseeseeeteees 14. 
Cymes in axils of 3-4 upper whorls only. Peduncles and pedicels 
with thick, patent hairs «0.00.0... cece 20. G. elbrussicum Pobed. 
Cymes in axils of almost all leaf whorls ..............:cscesseeseeseeees 15. 
Stem retrorsely scabrous. Each leaf whorl with rudimentary cymes 
along with fully developed ones. Pedicels with clusters of long, 
white hairs, exceeding flowers .............. 18. G. schischkinii Pobed. 
Stem not retrorsely scabrous. Each leaf whorl with normally 
developed cymes. Pedicels without clusters of white long hairs ... 
Beek IRE celles cere clinton Sou We, BNE AAT, MEI EO 16. 


Stem glabrous, with soft pubescence only on lower internodes. 
Leaves serrate along margin and midrib beneath ................ceseeee 
Sts Bo Pa seal Re URN oak ees Rae oth gS a 17. G. vernum Scop. 
Entire plant completely glabrous or ciliate at base of leaves ...... 
anil ean daek sae ROR ae tRacete 25. G. coronatum Sibth. and Sm. 
Plantimoreomless pubescentyd es A. Ae ee ehecsieeetsense 18. 
Stem mostly glabrous, base with minute scale-leaves, upper part 
pubescent, with closely-packed leaf whorls ..............:ccseseesee 
figs GRID: SRO eben 5k Att da BUR SOUR 27. G. sevanensis Pobed. 
Entire stem pubescent with patent hairs and scattered whorls of 
SreeniMeavesthint sieht ewe rete Mien see NERY SUC NS. aca 19. 
Cymes shorter than leaves. Mericarps glabrous. Leaf whorls 
SC Aten ep ei oe ae ae ccded cust 21. G. cruciata (L.) Scop. 


276 


291 


23(9).. 


26. 


2 


Cymes longer than leaves. Mericarps pubescent (rarely glabrous). 


Leaf whorls more or less COMPAaCt ............::cescccessccceeeecceseeeeeees 20. 
Leaves minute, 2—6 mm long, 1-3 mm broad ................:.0:0008 21. 
Leaves large, 5-21 mm long, 2.5—14 mm broad ....................- 22: 


Leaves 4—6 mm long, 2—3 mm broad, Leaf whorls so close together 
that leaves cover entire stem. Fruits glabrous .................:ceeeeee 
Se ee en A ee | eee oe 23. G. sosnowskyi Manden. 
Leaves 2—3 mm long, 1-1.5 mm broad. Leaf whorls close only 
BHDASE oc pe ees ie es ae 24. G. braunii Zelen. 
Leaves (5)7(10) mm long, (2.5)4(6) mm broad, elliptic or obovate, 
leaf whorls compact in upper half of stem. Entire plant densely, 
patently long-hairy, or leaves ciliate along margin and beneath 
along midrib, for rest glabrous. Fruits densely short-pubescent or 
PlabTOUSs., Sassi Bat 2 22. G. tauricum (Willd.) Roem and Schult. 
Leaves larger, (8)13(21) mm long, 8-14 mm broad, obovate, 
broadly elliptic or almost orbicular, leaf whorls somewhat spaced. 
Entire plant softly pubescent. Fruits with scattered, more or less 
long hairs or glabrous ...................05 26. G. kopetdaghense Pobed. 
Cymes furcate, terminal and in axils of upper leaves ............. 24. 
Inflorescence paniculate or cymes simple, 3-flowered terminal on 
stems and branches and in axils of upper leaves .................... 26. 
Leaves with one, distinctly prominent vein, lower leaves paired, 
cauline leaves in whorls of 4, 2 of which considerably larger than 
Othentwojtics.2iacked..senn.. beers 31. G. paradoxum Maxim. 
Leaves with 3 veins, leaves in whorls of 4 equal .................. 25. 
Leaves broadly elliptic, broadly ovate or almost orbicular, 
(9)15(21) mm long, (5)9(10) mm broad. Corolla lobes ovate 
hi aseahaet ania sais atc ca thad thea eet hn laatacanane 32. G. scabrum L. 
Leaves broadly obovate or orbicular, (14)15—20(25) mm long, 
(11)12-—15(17) mm broad. Corolla lobes ovate-lanceolate .......... 
mm, aaah ae ying. eomenaeding Bo2 33. G. kamtschaticum Steller. 
Cymes terminal and in axils of upper leaves, simple, 3-flowered 


Inflorescence terminal on stem and branches, paniculate ....... 30. 
Stem glabrous. Corolla 4-fid. Stamens 4 .0...... cece seeseeeeeeee 
aniets: Mensa S TRONS: 30. G. pseudopolycarpon Somm. and Lev. 
Stems retrorsely scabrous. Corolla 3-fid. Stamens 3. ............. Phe 
Leaves broad, large, (10)15—17(22) mm long, (1.5)4.5 (6.5) mm 
broad. Stem glabrous or weakly retrorsely scabrous, 15-50 cm 
tallyPlantperennial. 2ester: .itiee. pesos 39. G. trifidum L. 
Leaves narrower and smaller, (5)10(15) mm long, (1)1.5(2.5) 
mm broad. Stem conspicuously scabrous. Annuals ................ 29. 


DoD 


29. 


30. 


Sil 


32" 


277 


Pedicels short, (1.5)3(8) mm long. Leaves elliptic. Plants smaller, 
SDM erate nA BIRO. ONE 41. G. baicalense Pobed. 
Pedicels longer, (4)8—10(15) mm long. Leaves narrower. Plant 
larger, 10-30 cm tall ............ sees 40. G. ruprechtii Pobed. 
Leaves with one vein. Fruit surface granulate. Stems weaker, 
creeping, retrorsely scabrous. (Section Aparinoides Jord.) ...... Silt 
Leaves with 3, rarely with one vein. fruits glabrous, rugose or with 
dense hooked hairs. Stems glabrous or softly pubescent, straight, 
strong. (Section Platygalium DC.) ...............sssssseeseeseseseeeteteees 33. 
Stem strongly retrorsely scabrous along ribs. Leaves broad, almost 
rounded at apex, slightly revoluted along margin, densely retrorsely 
spinulose along margin and midrib beneath. Plants smooth ....... 
oe A RCE, SANTOS) I, ESE: 36. G. karakulense Pobed. 
Stem glabrous or scarcely scabrous along ribs. Leaves glabrous or 
scarcely scabrous along margin. Fruits microgranulate-tuberculate 
peeeihentn gal NOR eg a osoncttnetniismownedoerechdie Mie LAL 32: 
Plant larger, 50-80 cm tall. Stem thick; terminal panicle many- 
flowered, lax. Mericarps oblong, with indistinctly granulate surface 
Daten ire bene lon Palin... tase deetatveundedeiet 35. G. saturejaefolium Trev. 
Plant smaller, 30-70 cm tall, with slender stem. Upper panicles 
weakly expressed, cymes few-flowered, terminal on stem and 
branches. Mericarps globose, microgranulate-tuberculate .......... 
EDR RRS RI 34. G. palustre L. 
Fruits glabrous, rugose when dry because of pericarp being separate 
Gina Ariat ORI 28 ABER ORE 34. 
Fruit with dense hooked hairs or glabrous, but not rugose in dry 
form. Pericarp closely fitted in Mmericarp ...............::ssseseeeeeerens a7. 
Leaves lanceolate, attenuated toward apex, acute, thin, soft. Panicles 
lax /Spreadinigi: RRIRE Ue. SOIR Le 54. G. volgense Pobed. 
Leaves lanceolate-ovate or elliptic, not attenuated toward apex, 
obtuse, rigid or almost coriaceous. Panicles dense, compact ... 35. 
Leaves and stem glabrous, with short hairs only at nodes (pubescent 
only in hybrids with G. septentrionale Roem. and Schult., then 
fruits with pubescence scattered, on only one side). Inflorescence 
with few flowers, more or less lax ...... 53. G. ussuriense Pobed. 
Leaves and stem more or less pubescent along ribs ............... 36. 
Leaves, broad, large, (2)5.5-6.5(11.5) cm long, (0.6)1—2(3.5) cm 
broad, lower leaves softly pubescent often above and beneath or 
only beneath, middle and and upper leaves scabrous due to setaceous 
hairs along margin and veins beneath. Inflorescence at onset of 
flowering compact, later ovate OF COFYMDOSE ............eeceeeeeseeeees 
LPL EIS PGAD is ATE ESI Biles 0 BED. zee 52. G. articulatum Lam. 


293 


278 


Sis 


38. 


39: 


40. 


41. 


42. 


43(1). 


+ 


Leaves narrower and smaller, (2.5)4—4.5(5) cm long, 0.8-1.5 cm 
broad, all glabrous above, scabrous beneath and along margin 
due to short scattered antrorse setae. Inflorescence more or less 


laine mene cigt: SRe ry leer, ot ed a it 53. G. rubioides L. 
Leaves narrow, linear, (15)35(40) mm long, (0.5)1.5-4(9) mm 
broad eadieuivenes, areitven Pircorscbertarie fee inonatere mtorr mines oc. 38. 


Leaves broad, lanceolate-linear, lanceolate, oblong-lanceolate, 
broadly elliptic (rarely linear), 12-45-55(70) mm long, (2)6—10(25) 
MAIO... .nccccsnnrdansannhero Mel Meee gare teoMOE gre bal AN, apogee hs 39: 
Leaves 15—20 mm long, 0.5—1.5 mm broad, with one vein. Corolla 
3-3.5 mm across, with ovate, subacute lobes. Fruits with thick 
short, erect, straight setae ............ 58. G. amblyophyllum Schrenk. 
Leaves (27)35(40) mm long, (3)4(9) mm broad, with 1-3 veins. 
Corolla 3—5 mm across, with elliptic, short-acuminate lobes. Fruits 
sparsely pubescent with hooked setae or glabrous ...................... 
ues, a lunbie, oe alread mec ay 59. G. turkestanicum Pobed. 
Panicle corymbose. Upper branches long ending into corymbs 
almost equaling the terminal corymb. Leaves rhombic-lanceolate. 
Pniitselabrousi2 .f8inal.aek. i. aN ee 62. G. valantioides M.B. 
Panicle pyramidal, more or less spreading or dense, narrow. Leaves 
ovate, lanceolate or lanceolate-linear, rarely linear. Fruits mostly 
pubescent, rarely «glabrous).s:celete-aeseesies ees 40. 
Stem densely pubescent. Panicles at terminal on stem and all 
lateral branches, almost to base. Leaves pubescent on both surfaces 
with short appressed setae .............. 60. G. mugodsharicum Pobed. 
Stem glabrous or scabrous along ribs or pubescent along edges 
only at lower and upper internodes ...............:esseeeseeeseeeeseeeeees 41. 
Leaves linear or linear-lanceolate, (15)20(30) mm long, (2)3— 
4(5) mm broad. Panicle narrow, elongated ....... 56. G. boreale L. 
Leaves broadly elliptic or oblong-lanceolate, large, 28-70 mm 
long ;:4—24 mm ‘broad... 2nesssic:5. taki ie eee 42. 
Fruits with straight setaceous hairs. Leaves coriaceous, obtuse, 
broadly elliptic, or oblong-lanceolate, (28)30-40(45) mm long, 
(6)10(25) mm broad .............::ccccssecceeseeee 57. G. amurense Pobed. 
Fruits with hooked hairs. Leaves not coriaceous, lanceolate or 
oblong-lanceolate, gradually narrowed toward apex, (30)45-—55(70) 
mm tong,(4)7—8(12):mm broad Gal thd. eee ites eg. o.esentbhee 
sth. detene tenaies. teats erent 61. G. septentrionale Roem. and Schult. 
Cymes axillary, lax, few-flowered. Annual herbs with weak, 
creeping, retrorsely scabrous and, hence, often climbing stems. 


(Section Asparine RsSchmi:) ee ee) | ee heneearien ees fS 3. 44. 
Inflorescence more or less paniculate, or cymes terminal and in 
axils of upper leaves, few-flowered ..............:ccssssssesesssceeseeee 53. 


44. Pedicels recurved or scarcely inclined downward .................+ 45. 


294 


52 


53(43). 
+ 

54, 

+ 

55. 


+ 


56. 


Pediel sistraighte hate ee ocelot cers eee ea 47. 


MR tSie ATOMS, AUIDERCULALE, <.i..ccnasesennannsandevno dion eeeaaiae. 5 03 46. 


Fruits covered with white hooked hairs having thickened base 
BLU iethetn ties Lath apa SCY Ne 5. G. linezevskyi Pobed. 


. Fruits large, 4-6 mm broad, brown, acutely tuberculate ........... 


Seetn Bee nt nomen aed, aieibercom 4. G. tricorne Stokes. 
Fruits tiny, 1 mm long, 1.5 mm broad, very finely granular- 
fuberculate 2 ER. 10. G. transcaucasicum Stapf. 


“Pedicelsilone hairy? tintin ae seston wand La... 48. 


PedicelS)shortermthicksnot hairy eect ae 2200S. AL. a. 


. Flowers whitish or yellowish green .............ccccccsssssssseeeseeeeees 49. 


Flowers; purplet. ve. tite Se) epee Sache LE 50. 


. Corolla lobes with long cusp exceeding length of lobes ....... 


in Tk TR res nee EI dk Ee ie 9. G. trichophorum Kar. and Kir. 
Corolla lobes short acuminate ................ 8. G. tenuissimum M.B. 
Fruits densely white hairy ................008 13. G. eldaricum Grossh. 
Fruits glabrous or with scattered, white hairs ...............:cscsesee 
Ai, SENSOR ae snsde sEhartils £5 gest vee ete | 12. G. decaisnei Boiss. 


. Stem with patent, rigid hairs. Plant S—10 cm tall, conspicuously 


divaricately branched ......... 11. G. chloroleucum Fisch. and Mey. 
Stem retrorsely scabrous, climbing. Plant 10—100 cm tall, branches 
LEWD, Leth eies Ci tick ae AEE a cheat Menthe BR eta, sakatus 52. 
Plant large in all parts. Stems 20-100 cm tall, hispid at nodes. 
Leaves (13)30(52) mm long, (1.5)2(6) mm broad. Fruits 2.5-3 
mm long, 3.5-6 mm broad, covered with hooked hairs on 
tubercles. Cymes simple, 2—3-flowered, rarely 5-flowered, often 
reduced to single flower. Flowers white ............. 6. G. aparine L. 
Plant smaller. Stem 10-50 cm tall, glabrous at nodes. Leaves 
(10)15—20(40) mm long, (1)1.5—2(3) mm broad. Fruits 1-1.5 mm 
long, 2-3 mm broad, covered with hooked hairs without tubercles 
or-rarely with tubercles (var. Vaillantii). Cymes branched, 6—9- 
flowered, more rarely simple 3-flowered or reduced to single 
flower. Flowers greenish yellow ...............:0000 7. G. spurium L. 
Fruitwith granulateisurfacesc:! «227th So aaseee, -cscasteaes. nna ek 54. 
Fruits glabrous, smooth, or rugose, or pubescent ................00++ 63. 
Sténisismooths. 0.64. cReR ete bee Boe tino ee nda 55. 
Stems retrorseélyscabrous}eiei.eled. ots eich neat 60. 
Cymes more or less branched, terminal on stems and branches ... 
BE, URN NE ENING il cata cnnea cet acces saah Mapa Onan 56. 
Cymes simple, few-flowered (1-3), terminal on stems and branches 
or, in addition, axillary with solitary flowers .............:csccesee 59) 
eavesin whorls:efiGss beseties tenet. coe eles, ek a7. 
eaves) imiwhorls:of'5-6 cn .2os. nv) Ay ae... 58. 


295 


296 


280 
Si. 


+ 


58. 


59: 


60. 


Panicle almost corymbose, terminal on stems and branches. Fruits 
binate,.8. Gere BO RE 45. G. pumilum Murr. 
Cymes on peduncles, furcately or ternately branched, terminal on 
stems and branches. Fruits with 1 mericarp, rarely binate ......... 
fein le oe ono. kee pom 44. G. anisophyllum Vill. 
Leaves oblanceolate. Inflorescence terminal; cymes forming 
umbellate panicle. Pedicels short, equaling flowers or somewhat 
longer. Plant 5-30 cm tall ................. 42. G. hercynicum Weig. 
Leaves elliptic or oblong. Terminal inflorescence not conspicuous. 
Pedicels 2 times as long as flowers. Plants smaller, 5—10 cm tall 
ic Sew oe SR I IRE TEN 43. G. tianschanicum M. Pop. 
Cymes complex, doubly branched on long peduncles, greatly 
exceeding densely foliated vegetative shoots. Leaves narrowly 
linear or oblanceolate-linear, (4)10(15) mm long, 1—1.5 mm broad 
REP AMIN EHR Sib RL hs SEDI 46. G. vartani Grossh. 
Cymes simple, terminal— 3-flowered, and axillary— single- 
flowered. Peduncles and pedicels not longer than floral leaves. 
Leaves elliptic, 4-6 mm long, 1.5—2 mm broad ............:c:ceseeeeees 
donee, Metered fe .aed, bie BE 47. G. kiapazi Manden. 
Plant with extremely long, hairy peduncles ............... cesses 
See ONDA SS REO NI OME: POEL 49. G. dahuricum Turcz. 


+ Plant with peduncles lacking hairs .............cccssscseeseeseeereesees 61. 


63(53). 


65. 


. Leaves oblanceolate or lanceolate, tiny, (3)10(16) mm long, 


(DSC) unmubroddl 2S. Re AUR AS 48. G. uliginosum L. 
Leaves elliptic, oblong-elliptic or oblanceolate, large and broader, 
10-40 mm long, 2-14 mm broad 0.0.0... eee eeeeeseceeeeeseeeeeeeeees 62. 


. Leaves gradually tapered toward apex and abruptly short- 


acuminate, antrorsely spinulose along margin and vein beneath. 
Corolla 5-6 mm across. Fruits sparsely covered with lameller, 
white, lustrous haifs ................:sceeeeees 51. G. pojarkovae Pobed. 
Leaves rounded or mucronate at apex, retrorsely spinulose (rarely 
antrorsely) along margin and vein beneath. Corolla 3.5 mm across. 
Fruits with dense hairs ..................00 50. G. vassilczenkoi Pobed. 
Leaves linear-filiform, linear, more rarely oblong-linear or 
lanceolate-linear. Flowers golden-yellow, rarely cream or milk 
white. (Section Xanthogalium DC.) ..............cccessscesseeeseeeeees 64. 
Leaves lanceolate, oblanceolate, oblong-elliptic, rarely oblong- 
linear. Flowers white, rarely white-yellow. (Section Eugalium 
DG oe. cece la ee. RS TE ee 79. 


. Ovary and fruits glabrous ........... ccc eceeceeseseseesseeseeneenceneeneeeees 65. 


Ovary and fruits pubescent .............cccccccssssseesesessesseeseesneeneeseeees TS 
Pedicels black. Leaves reflexed after anthesis, short, broad ... 66. 
Pedicels yellow. Leaves not reflexed in fruit, long, narrow ... 67. 


297 


66. 


69. 


70. 


71. 


72. 


TSE 


74. 


281 


Leaves linear, 7-8 mm long, 1—-1.5 mm broad. Inflorescence 
spreading. Corolla milk-white ................ 71. G. achurense Grossh. 
Leaves oblanceolate, (10)20(25) mm long, (1.5)2.5(4) mm broad. 
Inflorescence oblong, narrow. Corolla golden-yellow ............... 
seats. «sala cepa eemaaers Mactan rennin hes aemrmetins sul 70. G. aureum Visiani 
Leaves oblong-linear. Stem often glabrous on lower internodes, 
elsewhere short-pubescent ...........sssscsssesesssssscssssssseseseseeseaeseaceees 68. 
Leaves lanceolate-linear, linear or filiform ................:eesseeeeee mi. 
Leaves long and narrow, (9)20—25(35) mm long, (0.5)1(1.5) mm 
broad, long cusp at apex. Corolla large, 4 mm across ................ 
De ee ee ee 67. G. anfractum Somm. and Lev. 
Leaves shorter and broader, (18)25(30) mm long, (1.5)2(3) mm 
broad, at top with short mucro. Corolla smaller, 3 mm across ... 69. 
Stems short-pubescent, glabrous on lower internodes, 4-angled. 
Leaves in whorls of 6-10 ................... 65. G. wirtgenii F. Schult. 
Stems glabrous, with ring of velvety hairs at base of leaf or 
pubescent, almost terete or terete only at base, above 4-angled, 
with glabrous ribs. Leaves in whorls of 8-10 ..............ccscccsseeeeees 70. 
Leaves large, (20)40(50) mm long, (0.5)1(2.5) mm broad. Panicle 
long, equaling 1/3 or 1/2 length of stem, spreading, corolla lobes 
subobtusewiede: ia geeattinat. ek 69. G. pamiroalaicum Pobed. 
Leaves (20)25(28) mm long, (1)2(3) mm broad. Panicle shorter, 
more or less narrow, oblong, not very dense. Corolla lobes 
mMUCTONAte HRB. veh. Geese ses. eee: 68. G. majmechense Bordz. 
Leaves lanceolate-linear, (6)7—8(15) mm long, (0.5)0.75—1.5(2) mm 
broad. Stems with conspicuous, nearly winged ribs. Panicle narrow, 
elongated, interrupted ........... 66. G. fistulosum Somm. and Lev. 
Leaves linear or filiform, (8)8—20(35) mm long, (0.25)1(1.5) mm 
broad. Stems 4-angled, without winged ribs. Panicle broad, more 
Omlessuspreadin Gyo. x csln.jcthedeeaess 8. heed erees beecobers, pelecouanels cd... a2. 
Panicle extremely lax; cymes ternately branched. Corolla lobes 
with long cusp (up to half length of lobes). Leaves filiform, 
(8)19(18) mm long, 0.25-O0.75 mm broad ..........cceceeseeeseceeeeeeees 
asteioaaap eoavetad oS Man ae aise, dba 64. G. subuliferum Somm. and Lev. 
Panicle dense, sometimes spreading; cymes doubly ternately 
branched. Corolla lobes subobtuse. Leaves linear or filiform, (10)18— 


20(35) mm long, (0.5)1(1.5) mm broad ............... 63. G. verum L. 
Stemeandyleaves. .plabrois) fcr lest con dedd. oie ae 74. 
Stemipand: leavesspubescenit esses exteie: sei deers roca TS: 


Panicle more or less spreading. Plants larger, 30-125 cm tall. Leaves 
usually with revolute margin .............:.cscceeeeee 63. G. verum L. 


298 


282 


78. 


Panicle very narrow, small, terminal stem. Plants smaller, 40-70 
cm tall. Leaves without revolute margins ...............:csseeeseeeeeeeees 
es eh aie Sad AER ENE 73. G. densiflorum Ldb. 


. Entire plant, grayish due to dense pubescence. Corolla lobes hairy 


outside. Panicle narrow, interrupted .................eceeeseseeeeceeeeeeeees 76. 
Plant not densely pubescent, not grayish. Corolla lobes glabrous 
outside. Panicle not interrupted, often spreading ..................... 77. 


. Leaves reflexed, tiny, (5)6(11) mm long, (0.5)9.75(1) mm broad. 


Plant small, 6—35 cm small .................:eeee 77. G. kutzingii Boiss. 
Leaves not reflexed, larger, 14-20 mm long, 0.5—1.5 mm broad. 
Plant larger, 30-50 cm tall ............. 76. G. antropatanum Grossh. 


. Panicle narrow, short, more or less lax. Leaves smaller (7)10(15) 


mm long, (0.5)0.75(2.5) mm broad. Plant smaller, 8-30 cm tall 
HOE, 75. G. saurense Litw. 
Panicle long, broad, more or less spreading, dense. Leaves larger, 
(20)35—45(55) mm long, (0.25)0.75(2.5) mm broad. Plant larger, 
257130 cintallts: BMS OR AR ee lea: 78. 
Leaves narrow, (20)25(30) mm long, (0.25)0.5(0.75) mm broad. 
Flowers light yellow. Eurasian plant ... 74. G. ruthenicum Willd. 
Leaves broader, (20)35—45(55)mm long, (0.5)0.75(2.5) mm broad. 
Flowers cream colored. Far Eastern plant ..................000 
SUERTE RON TRUE Fe RIE De 72. G. lacteum (Maxim.) Pobed. 


79(63). Entire plant gray due to dense pubescence of more or less long 


+ 


80. 


81. 


82. 


soft hairs. Fruits pubescent .............s:ecseeeee 84. G. caspicum Stev. 
Fruits glabrous or rarely pubescent only at base or over entire 
surface. Plant not gray due to pubescence, although stems or 
leaves)may Be pubescent ft? .20.01. 2, a 80. 
Leaves in whorls of 4—7, coriaceous, thick. Stems with woody 
caudex at/base an Ae a RR eR) 81. 
Leaves in whorls of 4 -6-8, not thick. Stems without caudices at 


Leaves in whorls of 4. Ovary and fruits hispid only at base or up 
to apex, or else completely glabrous. Stems retrorsely scabrous or 
lABPOUS AA a hee I BA 92. G. coriaceum Bge. 
Leaves in whorls of 6—7. Fruits microgranulate, covered with 
setulose tubercles. Stems asperate with very short, dense spinules, 
directed sideways ....... 93. G. brachyphyllum Roem anad Schult. 
Fruits globose, swollen, often with one mericarp, 3—3.5 mm in 
diameter, white, shining, with lax, serrate pericarp ........... 
LR I REN IN co BIE EIR, FE 90. G. bullatum Lipsky. 


Fruits not swollen, 1—-2.5 mm broad, brown ...........:cscceseeeeees 83. 
. Stems glabrous, rarely pubescent in lower part ............:.:0000 85. 
Stenis pubescent. 24.¥08 NON De. Pe, ae ood 84. 


84. 


85. 


86. 


87. 


88. 


299 91. 


283 


Stem and leaves on both surfaces velutinous. Panicle narrow, 
terminal, fairly long, but not extending to middle of stem ...... 
PA ISIN Oi Ce. Meme 5 Ln hed See 85. G. grusinum Trautv. 
Stem and leaves with thick, more or less long, patent hairs. Panicle 
very short, corymbose, small-flowered ..............cssceesseeeseeeeeeoees 
SEED NG ro Mahe he sh 91. G. grossheimii Pobed. 
Leaves intwhorls Git4=5=6' }...2ebacewl bas. metus. wee... 86. 
Beavessin whorlsioM6=B) siti set el eek eA 88. 
Stems 5—35 cm tall, herbaceous. Leaves linear, short, (5)7—11(20) 
mm long, (0.75)1—2(2.5) mm broad. Corolla lobes mucronate ... 
SaaS Rika S CONOR es AE DTA at a, A ala tinteed eb levis Adeeb 87. 
Stems 30-70 cm tall, woody at base. Leaves linear, 22-40 mm 
long, 0.5—4 mm broad. Corolla lobes with or without a long cusp 


pea Mill git CM. OE LIS Onel e 195 <A 3 OLE aoe, eh pee Fee 88. 
Leaves linear, in whorls of 6, (5)11(20) mm long, (0.75)1(1.5) mm 
broad. Anthers yellow «....2.2.2...2.0cc000 86. G. hyrcanicum C.A.M. 


Leaves oblanceolate, in whorls of 4-5, upper leaves in whorls of 5— 
6, (5)7—10(11) mm long, (1)1.5—2(2.5) mm broad. Anthers black 
a ae Doak ROD, Leben Nace atta 8. G. czerepanovii Pobed. 
Angles of stem obtuse, with broad, white hairs. Leaves narrowly 
linear, (22)(25)(30) mm long, (0.5)1(1.5) mm borad. Corolla lobes 
gradually attenuated into elongated cusp. Flowers white ............. 
RE Se Rees. Aeneas 88. G. mite Boiss. and Hohenack. 
Angles of stem with narrow, white bands, acute. Leaves (25)30 
(40) mm long, (1.5)2(4) mm broad. Corolla lobes without or with 
minute mucro. Flowers flesh colored or pink ............. cesses 
PEELS RROD eR ee 89. G. leiophyllum Boiss and Hohenack. 
Plants with ascending, straight, mostly glabrous stem. Peduncles 


and pedicels more or less thick, ‘SHOMt 25....¢2-0056¢4-.-2-cessocesusesoeoenee 90. 
Plants tall, with nutant, glabrous, lustrous stems and trichoid, long 
pedunelesvandapedivels MAaee. Beco alee I Zs eke S3. 
Leaves more or less long, usually 15-20 mm long and 1.5—2.5 mm 
broad, soft. Flowers! whites...) enw nat ie 91. 
Leaves more or less short, usually 6-15 mm long, 1.5—2 mm _ broad, 
Stilt pPlowenrsreraymyellowi tan vie: Folie ees ied late) Nea 92. 


Panicle spreading, with long, divaricate branches. Peduncles 
divaricate at time of anthesis. Leaves linear-oblong or oblanceolate, 
(10)20(25)mm long, (1.5)2.5(4) mm broad ... 78. G. mollugo L. 
Panicle narrow, with short erect branches. Peduncles straight at 
time of blossoming. Leaves linear-lanceolate, (9)15(18) mm long, 
GIS =2@) mimi ibroaG? ne ale 79. G. erectum Huds. 


300 


284 


92. Panicle narrow, long or somewhat spreading. Leaf whorls distant, 
leaves (9)10—15(18) mm long, (1)1.5(3) mm broad. Corolla lobes 
oblong, with long cusp at tip, not incurved. Corolla 3.5—-4 mm 
across. Stamens with long filament: «2.0.0.0... eteeeeteteeeeeees 
ep coc 81. G. calcareum (Alb.) Pobed. 

+ Panicle narrow, very short, terminal. Leaves in compact whorls. 
Leaves shorter and broader, (5)6—7(12) mm long, (1)1.5—2(2.5) 
mm broad. Corolla lobes ovate, with incurved, short mucro ....... 
TLR). ROM LIBROS SOURS | PINION 80. G. juzepcezukii Pobed. 

93. Leaves large, (23)40—45(60) mm long, (3.5)5—7(9) mm broad, green 
above; bluishibene ath 3) c..cccscccose Meateoot 82. G. schultesii Vest. 

+ Leaves smaller and narrower, 22—23 mm long, 1.5—3 mm broad, 
green above, paler beneath, not turning bluish ..........0 0... eee 
wiaciscrad otcaiutis tbe otic ahs coe eee ie eae ee Dea 83. G. polonicum Blocki. 


Section 1. DEPAUPERATA Pobed. sect. n. in Addenda XXII, 709.— 
Trichogalia DC. Prodr. IV (1830) 599 p.p. pro subsect.—Cymes 
axillary, rarely terminal, reduced to one flower or 2—3 flowers, on 
long peduncles. Leaves in whorls of 4 unequal—2 smaller than the 
other two, decussate, cauline leaves in whorls of 6 equal, with 1 vein, 
short acuminate. Fruits covered with long hooked hairs. 

Besides our species, this section includes G. pauciflorum Bge. 
from northern China and the Himalayas and G. handelii Cuf. from 
northern China. 


Subsection 1. Quadrifolia Pobed. subsect n. in Addenda 
XXII, 710.—Cymes axillary, reduced to one flower. All leaves in 
whorls of 4, 2 smaller than the other two, decussate. 


1. G. soongoricum Schrenk in Fisch. et Mey. Enum. pl. nov. I 
(1841) 57; Kar. and Kir. in Bull. Soc. Nat. Mosc. XV, 372 (ex scrip. 
103); Ldb. Fl. Ross. II, 421; O. and B. Fedch. Konsp. Fl. Turkest. 3, 
141; Kryl. Fl. Zap. Sib. X, 2591. 

Annual. Root slender, filiform, reddish or brown. Stem weak, 
lodged, slender, branched, 5-30 cm tall, glabrous, rarely (in specimens 
from Ashkhabad) with scattered , long hairs, 4-angled. Leaves in whorls 
of 4, 2 smaller than the two, decussate, elliptic, (7)12-—15(23) mm 
long, (2)5(8) mm broad, short-acuminate often with scarcely distinct 
mucro, with short petioles, singled-veined, thin, glabrous on both 
surfaces, rarely (in specimens from Ashkhabad) covered with scattered 
erect hairs. Cymes axillary, 1-flowered, on glabrous (12)20(40) mm 
long pedicels, longer than leaves, occasionally with a pair of tiny 
bracts, somewhat higher than the middle of pedicels; rarely 2-3 flowers 


303 


285 


over total length of peduncle on short pedicles, usually horizontally 
directed. Corolla white, 0.5—1 mm across, with 4, rarely 3 (var. trifidum 
Kryl.), broad, ovate-deltoid, acute lobes. Stamens corresponding to 
lobes 4, rarely 5, Anthers yellow. Style bipartite almost from middle. 
Fruits binate, rarely with one mericarp, the second rudimentary, 2 mm 
long, 2.5 mm broad, covered by long hooked white hairs. June to 
August (Plate XVI, fig. 1). 

Moist spruce and juniper forests, in subalpine and alpine mountain 
zones.— Western Siberia: Altai. Soviet Central Asia: Dzhungaria- 
Tarbagatai, mountainous Turkmenia, Pamiro-Alai, Tien Shan. Endemic. 
Described from foothills of Dzhungarian Ala Tau, Baskan River. Type 
in Leningrad. 

Note. Plants collected by D.I. Litvinov near Ashkhabad in the 
Karanki gorge at an altitude of about 3000 m differ from G. 
soongoricum in having pubescent stems and leaves with scattered, 
white hairs; a long white mucro at the apex of the upper leaves and 
bracts; and shorter pedicels. It is possible that these plants represent 
a separate race, close to G. soongoricum, which, for lack of material, 
is difficult to establish. It should be noted that the Kopet-Dag specimens, 
closest to the Ashkhabad specimens do not differ from G. soongoricum. 

Regel described var. diffusum Rgl. (O. and B. Fedch. Konsp. fl. 
Turkest. 3 (1909) 141) from the alpine zone of Tien Shan with very 
small leaves (4-5 mm long, 1—1.5 mm broad) and tiny mericarps 0.75 
mm long, 1.25 mm broad. 


Subsection 2. Senifolia Pobed. subsect. n. in Addenda XXII, 
710.—Cymes axillary or terminal. Cauline leaves in whorls of 6, basal 
and upper leaves in fours, all equal. 


2. G. triflorum Michx. Fl. Bor. Am. I (1803) 80; DC. Prodr. IV, 
601; Ldb. Fl. Ross. II, 413; Schmalh. Fl. II, 15, Kom. and Alis. Opred. 
rast. Dalnevost. kr. II, 961; Kryl. Fl. Zap. Sib. X, 2583.—G. suaveolens 
Wahlenb. Fl. lapp. (1812) 48.—G. triflorum var. viridiflorum DC. l.c. 
(1830) 600.—G. bryophilum Goldbach apud Weinm. Elenchus pl. h. 
Pawlowsk (1824) 69.—Ic.: Syreistsch. Fl. Mosk. gub. III, 194; Fedch. 
and Fler. Fl. Evrop. Ross. 913, fig. 899.—Exs.: Dorfler, Herb. norm. 
No. 5078; Pl. Finl. exs. No. 362 a and b; GRF, No. 267. 

Perennial. Rootstock slender, creeping. Stems prostrate, weak, 25- 
60 cm tall, 4-angled, with slender retrorse hairs along ribs, nodes 
usually thicker, branches several. Lower and upper leaves in whorls 
of 4, middle in 6, oblong-lanceolate, (15)20—25(40) mm long, (6)7- 
8(15) mm broad, short-acuminate, gradually attenuated at base into 
short petiole or almost sessile, single-veined, glabrous or diffusely 


286 


4 
| Be Ns ’ 
7  , i y 

j A> 

D o ny, 

E q 

f 
WwW u 


aN bins 
iN i 
i\ 74 AN Prt AY LEE 
ay | K\ A\K i y Z 
ae sis ‘\ VANES 
Ji 3 i << } 
QY Nay 


EIN 


‘ \ ; 


Plate XVI. 


301 1. Galium soongoricum Schrenk., general appearance, corolla, ovary with styles, 
fruit—2. G. tauricum (Willd.) Roem. and Schult., general appearance, corolla, 
ovary with styles, fruit—3. G. kopetdaghense Pobed., general appearance of 
plant, corolla, ovary with styles, fruit—4. G. krylovii Iljin, general appearance 

of plant, whorl of leaves from below. 


304, 


287 


antrorsely hairy on upper surface toward tip, retrorsely spinulose along 
vein beneath. Floral leaves subulate-lanceolate, 4-5 mm long, 0.25- 
0.5 mm broad. Cymes axillary, on long, glabrous peduncles with 3 
flowers on shorter similarly glabrous pedicels. Corolla white (in 
European plants) and green (in Siberian plants— var. viridiflorum DC.), 
3.5—4 mm across, with lanceolate, gradually long-acuminate lobes. 
Anthers yellow. Style bipartite in upper third. Fruits binate, often with 
one, almost globose mericarp, 1.5 mm long, 2 mm broad, covered 
with long, white, lustrous hooked hairs. June to August. 

In shady conifers, mixed coniferous broad-leaved and birch forests, 
in wet places with moss cover.— European USSR: Karelia-Lapland, 
Dvina-Pechora, Baltic Region, Ladoga-IImen, Upper Volga, Volga- 
Kama, Volga-Don (only in northern part). Western Siberia: Ob’, Irtysh. 
Eastern Siberia: Yenisei (very rarely), Lena-Kolyma (only along Aldan 
River), Angara-Sayan, Dauria. Soviet Far East: Kamchatka, Uda River 
area, Ussuri, Sakhalin. General distribution: Scandinavia, Japan, China, 
North America. Described from Canada. Type in Paris. 

Note. As to the report of this species in the Himalayas (Kry]l. FI. 
Zap. Sib. X, 2584), it is in all probability erroneous. The single 
specimen from Kashmir available to us, resembling G. triflorum, differs 
from it in having very long and narrow leaves. 


3. G. trifloriforme Kom. in Tr. SPb. bot. sada XVIII (1901) 428 
and XXV (1908) 497; Kom. and Alis. Opred. rast. Dalnevost. kr. II, 
961.—Ic.: Kom. l.c. XXV, tabl. IX. 

Perennial. Rootstock very slender, creeping, producing 
subterranean shoots. Stem weak, ascending, straight, 30-40 mm tall, 
branches few, with slender scattered retrorse hairs along ribs, or almost 
glabrous. Lowermost leaves paired, lower ones in whorls of 4, upper 
and middle in 6, oblong-obovate or obovate, (18)25(28) mm long, 
(5)7(10) mm broad, with short, flat mucro at apex, gradually narrowed 
at base into short petiole or almost sessile, antrorsely appressed, hairy 
on upper surface toward apex with short sparse, retrorse appressed 
hairs along middle vein beneath; floral leaves paired, lanceolate, small, 
(2)4(5) mm long, (0.5)1.5(3) mm broad. Cymes terminal and in axils 
of upper leaf-whorl with 2-3 divaricate pedicels, many times as long 
as flowers. Peduncles longer than leaves, together with pedicels, 
glabrous. Corolla 3 mm across, with broad lanceolate, long acuminate 
lobes. Stamens on long filaments. Anthers dark. Style long, highly 
exserted from flower, bipartite from middle. Fruits 1 mm long, 2 mm 
broad, binate, often with 1 mericarp, covered with long (longer than 
foregoing species), white, lustrous curved hairs. June to August. 


305 


288 


In moss-covered, mixed and coniferous forests on mountain slopes 
and in river valleys, in wet soil.—Soviet Far East: Ussuri, Sakhalin. 
General distribution: Japan, Korea, China. Described from Korea. Type 
in Leningrad. 


Section 2. APARINE DC. Prodr. IV (1830) 605, p.p., K. Schm. in 
Pflanzenfam. IV, 4 (1897) 152.—Euaparine Lange ex Willk. and 
Lange, Prodr. fl. Hisp. 2 (1870) 325; Rouy and Camus, FI. Fr. VIII, 
48.—Cymes axillary, 3—5-flowered, more rarely reduced to one flower, 
on more or less long peduncles. Leaves in whorls of 6-12, all equal, 
single-veined, cuspidate. Fruits glabrous or pubescent, erect at tip or 
curved at base, but not hidden under leaves. Annuals, often clinging 
herbs with weak retrorsely scabrous stems. 


Subsection 1. Camptopoda Boiss. FI. or. II (1875) 67.— 
Camptaparine K. Schm. in Pflanzenfam. IV, 4 (1897) 152 (subsect.).— 
Fruiting pedicels recurved. Corolla cream colored. 

This subsection also includes G. saccharatum All, besides our 
species, which is closely related to G. tricorne. G. saccharatum All. 
is widespread in Central Europe and in the Mediterranean Region from 
the Canary Islands to Palestine. 


4. G. tricorne Stokes in With. Bot. Arr. Brit. Pl. ed. II, 1 (1787) 
153; DC. Prodr. IV, 608; Ldb. Fl. Ross. II, 419; Boiss. Fl. or. III, 67; 
Schmalh. Fl. II, 12; O. and B. Fedch. Perech. rast. Turkest. III, 141; 
Grossh. Fl. Kavk. IV, 29.—Valantia tricornis M.B. FI. taur.-cauc. II 
(1808) 438 and III (1819) 640.—V. spuria Pers. Syn. I (1805) 129.— 
V. triflora Lam. FI. Fr. 3 (1778) 384 (excl. Syn.).—Ic.: Malkov. Tr. 
po. prikl. bot. gen. i. sel. XXV, IV (1931) 158; Hegi, Ill. Fl. VI, 1, 
226, f. 122 k-m.—Exs.: Fl. Palaest. exs. No. 185. 

Annual. Stems decumbent, simple, 4-angled, 10-60 mm tall, 
clinging due to long, acute, retrorse spinules along ribs. Leaves in 
distant whorls of 6—8, linear-lanceolate, (8)20(30) mm long, (1.5)2.5(4) 
mm broad, cuspidate, gradually attenuated at base, single-veined, with 
clinging retrorse spinules along margin and vein beneath; floral leaves 
paired, unequal, lanceolate, (4)5(7) mm long, (0.5)1(1.5) mm broad. 
Cymes axillary, paired, 3-flowered, not exceeding leaves, peduncles 
and pedicels thick, 4-angled, covered with spinules along ribs, recurved 
in fruit. Flowers bisexual, rarely cymes with only male flowers. Corolla 
cream colored, 2.5—3 mm across, with lanceolate lobes. Anthers yellow. 
Style bipartite from base. Fruits binate, 4 mm long, 6 mm broad. 
Mericarps brown, globose, densely acutely tuberculate, very often | 
mericarp not fully developed. April to August. 


306 


289 


In fields as weed, along roads, rarely on rubbly steppe slopes.— 
European USSR: Upper Volga, Black Sea Region, Crimea. Caucasus: 
All regions. Soviet Central Asia: Kara Kum, mountainous Turkmenia, 
Syr Darya, Pamiro-Alai, Tien Shan. General distribution: Central and 
Atlantic Europe, western and eastern Mediterranean, Balkans-Asia 
Minor, Armenia-Kurdistan, Iran-Afghanistan, North America 
(introduced). Described from England, where it occurs as an adventive. 
Type in London. 


5. G. linczevskyi Pobed. sp. n. in Addenda XXII, 710. 

Annual. Stem straight, 3-7 cm tall, simple, 4-angled, with long, 
white soft, patent hairs along angles in lower internodes, retrorsely 
scabrous due to short spinules elsewhere. Leaves in whorls of 6, 
uppermost in whorls of 2-4, oblanceolate or oblong-oblanceolate, 10— 
13 mm long, (1.5)3(4) mm broad, abruptly short-acuminate at apex, 
gradually narrowed at base into more or less elongated petiole, single- 
veined, tomentose on both surfaces, especially in young leaves with 
long, white, thin retrorse spinules along margin and beneath at base of 
vein, or without tomentose pubescence (specimens from Zeravshan 
Range). Cymes single-flowered in axils of leaves, on slender retrorsely 
scabrous pedicels, longer than leaves, slightly curved in middle and 
with one, 1.5—3 mm long, 0.5—1 mm broad, bract. Corolla much smaller 
than ovary, 1.5 mm across with ovate, acute lobes. Anthers yellow. 
Style bipartite somewhat above base. Fruits binate, rarely with one 
mericarp, 2.5 mm long, 4 mm broad; mericarps subglobose, covered 
with white thick hairs having thickened base and hooked tip, with or 
without tomentose pubescence covering the fruit. June to July. 

In upper mountain belt in shady screes and rocks, in juniper 
stands.— Soviet Central Asia: Pamiro-Alai. Endemic. Described from 
Babatag Range. Type in Leningrad. 


Subsection 2. Leucaparinea Boiss. FI. or. III (1875) 68.— 
Euaparine Lange in Will. and Lange, Prodr. fl. Hisp. II (1870) 325 
(pro sect.).—Fruiting pedicels straight, flowers white. 


6. G. aparine L. Sp. pl. (1753) 108; M.B. Fl. taur.-cauc. I, 105 
and III, 110; DC. Prodr. IV, 608; Ldb. FI. alt. I, 133 and Fl. Ross. II, 
419; Boiss. Fl. or. III, 68; Schmalh. Fl. II, 13, p.p.; Grossh. Fl. Kavk. 
IV, 29.—Aparine hispida Moench. Meth. pl. (1794) 640.—Ic.: Fedch. 
and Fler. Fl. fig. 896 (under the name G. spurium L.); Hegi, Ill. Fl. VI, 
1, tab. 249, f. 1.—Exs.: Fl. Gal. and Germ. exs. No. 3408; Fl. exs. 
austro-hung. No. 2226; Pl. Finl. exs. No. 954 (sub G. vaillantii DC.); 
Fl. Pol. exs. No. 830. 


307 


290 


Annual. Stem weak, 30-100 cm tall, clinging, 4-angled, retrorsely 
spinulose along ribs, somewhat thickened and often hispid at nodes. 
Leaves in whorls of 6-8, linear-lanceolate, (13)30(52) mm long, 
(1.5)2(6) mm broad, abruptly short-acuminate at apex, gradually 
narrowed at base, sessile, retrorsely spinulose along margin and vein 
beneath, upper surface glabrous or with short, antrorsely appressed 
rigid hairs; floral leaves smaller, (4)7(9) mm long, (0.75)1(1.5) mm 
broad, in whorls of 6-8 at base of peduncle and 1-2 below pedicels. 
Peduncles axillary, long, scabrous due to spinules; pedicels more or 
less short, glabrous or slightly scabrous, straight in fruit. Cymes simple, 
2-3 flowered, rarely 5-flowered, often reduced to single flowers. 
Flowers bisexual. Corolla white, 1.5 mm across, with oblong lobes. 
Anthers yellow. Style bipartite almost up to base. Fruits binate, 2.5— 
3 mm long, 3.5—6 mm broad, covered with hooked hairs borne on 
tubercles. June to September. 

Fields, gardens, railroad embankments, waste places as weed.— 
European USSR: All regions. Caucasus: All regions. Western Siberia: 
Ob’, Upper Tobol, Irtysh (very rarely). Eastern Siberia: Yenisei, Lena 
Kolyma, Angara-Sayan, Dauria (rarely, introduced). Soviet Far East: 
Ussuri (rarely, introduced). Soviet Central Asia: mountainous 
Turkmenia, Syr Darya, Pamiro-Alai, Tien Shan (introduced). General 
distribution: Scandinavia, Central and Atlantic Europe, western and 
eastern Mediterranean, Balkans-Asia Minor, North America. Described 
from Europe. Type in London. 

Note. In the mountains of Soviet Central Asia [Nuratau, Talass 
Ala Tau, Chatkal range, Kopet Dag (Mt. Saandak)] there is a form of 
this species with short and broad, thin leaves, which often has been 
identified by botanists as G. tenerum Schl. or as close to it. Apart 
from the size and shape of the leaves, this form is in no way different 
from G. aparine L. 


7. G. spurium L. Sp. pl. (1753) 106; DC. Prodr. IV, 608; Ldb. FI. 
alt. I, 134; Kar. and Kir. in Bull. Soc. Nat. Mosc. XIV, 3 (1841) 433; 
Grossh. Fl. Kavk. IV, 29.—G. vaillantii DC. Fl. Fr. 4 (1805) 1; DC. 
Prodr. IV, 608; Grossh. Fl. Kavk. IV, 29.—G. aparine Y. spurium 
Koch. Synop. (1837) 330; Ldb. Fl. Ross. II, 420.—G. aparine B. 
vaillantii Koch, |.c. 330; Ldb. l.c. II, 420; Boiss. FI. or. III, 69; Schmalh. 
Fl. II, 13.—G. spurium L. var. vaillantii Gren. and Godr. Fl. Fr. II 
(1850) 44.—Koch in Linnaea, XVII (1843) 33; Ldb. l.c. 420.—G. 
agreste &. echinospermum Wallr. sched. 59; Rupr. Fl. ingr. 494.—Ic.: 
W. and K. l.c. tab. 202; Hegi, Ill. Fl. VI, 1, f. 123 e, f. 

Annual. Stem 10-50 cm tall, usually glabrous at nodes. Leaves 
oblanceolate or linear-lanceolate, (10)15—20(40) mm long, (1)1.5—2(3) 


308 


291 


mm broad, with base gradually attenuated into more or less long petiole, 
with thin long cusp at apex. Cymes axillary, ternately or furcately 
branched, 6—9-flowered, rarely simple, 3-flowered or reduced to single 
flower. Flowers greenish yellow. Fruits 1-1.5 mm long, 2-3 mm broad, 
densely pubescent with curved hairs, without tubercles (or rarely with 
tubercles) at base or glabrous, otherwise not different from earlier 
species. April to August. 

In scrub, coastal sands, crops, gardens as weed.— European USSR: 
Karelia-Lapland, Dvina-Pechora, Ladoga-Ilmen, Baltic Region, Upper 
Volga, Volga-Kama, Upper Dnieper, Middle Dnieper, Volga-Don, 
Upper Dniester, Black Sea Region, Crimea, Lower Volga. Caucasus: 
All regions. Western Siberia: All regions. Eastern Siberia: All regions. 
Soviet Far East: Kamchatka, Ussuri. Soviet Central Asia: Aralo- 
Caspian, Balkhash Region, Dzhungaria-Tarbagatai, Kyzyl-Kum, Kara- 
Kum, mountainous Turkmenia, Syr Darya, Pamiro-Alai, Tien Shan. 
General distribution: Central Europe, Mediterranean, Balkans-Asia 
Minor, Iran-Afghanistan, India-Himalayas, Mongolia, North America. 
Described from France. Type in Paris. 

Note. For this species we are retaining the Linnaean name, 
following the rules of nomenclature. Under this name Linnaeus 
described the form with glabrous fruits that occurs sporadically in 
different parts of the range of the widely distributed G. vaillantii DC., 
which has pubescent fruits and was described after Linnaeus. In relation 
to G. aparine L., this is a corresponding species. In Asia, G. aparine 
occurs only as an introduced plant, but G. spurium L. (G. vaillantii 
DC.), clearly predominating in Asia, is also fairly widely distributed in 
Europe, within the range of G. aparine. G. spurium L., having such a 
wide geographical range, is not morphologically homogeneous: Siberian 
plants differ considerably in being taller, with long, narrow leaves, but 
the pubescence of their fruits without tubercles and the absence of 
pubescence at the nodes of the stem are unmistakable characters of G. 
spurium. Caucasian plants of this species are small plants with small 
leaves and fruits, occasionally with tubercles at the base of the hairs 
covering them; the stem usually has glabrous nodes. 


Section 3. PSEUDOAPARINE Lange in Willk. and Lange. Prodr. fl. 
Hisp. II (1870) 322.—Xanthaparinea Boiss. Fl. or. III (1875) 72, pro 
subsect.—Cymes axillary, doubly-triply furcate, ternate or simple with 
3 flowers. Leaves in whorls of 4—6—8, often unequal, very narrow, 
linear, single-veined, cuspidate. Fruit glabrous, rarely pubescent, 
peduncles longer than leaves, along with trichoid pedicels. Annuals, 
very thin, tender herbs with glabrous, rarely weakly retrorsely scabrous 
stem. 


309 


292 


Series 1. Tenuissima Pobed.—Flowers yellowish. 
Besides our species, this series includes G. divaricatum Lam. from. 
the western Mediterranean. 


8. G. tenuissimum M.B. FI. taur.-cauc. I (1808) 104 and III (1819) 
107; DC. Prodr. IV, 607; Ldb. Fl. Ross. II, 418; Boiss. Fl. or. III, 73; 
Schmalh. Fl. II, 13; Grossh. Fl. Kavk. IV, 29.—Exs.: GRF, No. 268; 
Fl. exs. austro.-hung. No. 2227. 

Annual. Stem weak, slender, ascending, 30-50 cm tall, strongly, 
repeatedly divaricately branched from base, 4-angled, weakly expressed 
ribs in upper part, ribs scabrous due to short retrorse setae, often 
distributed in lower internodes in 2 rows. Leaves in whorls of 4-8, 
linear-lanceolate, (4)8(11) mm long, 0.5—-1 mm broad, apex gradually 
long-acuminate, ending into a short cusp, gradually narrowed at base 
into short petiole or subsessile, single-veined, antrorsely setose, and 
hence scabrous along margin and vein beneath; floral leaves in whorls 
of 4-6, uppermost identical, (1)2(2.5) mm long, 0.15—0.25 mm broad, 
with pubescence similar to that of stems. Cymes numerous in axils of 
leaf whorls of main stem and branches. Peduncles doubly-triply furcate 
or ternate, together with trichoid pedicels. Pedicels (5)7-10(17) mm 
long, glabrous, clavate at top. Peduncles longer than leaves diffusely 
setulose. Corolla whitish or yellowish green, 1.5—2 mm across, with 
oblong-elliptic shortly acuminate lobes, with incurved cusp. Anthers 
yellow, on short filament, shorter than lobes. Style bipartite, almost 
from the very base. Fruits binate, 1 mm long, 1.25 mm broad. Mericarps 
falcate, just about oblong, glabrous, very finely tuberculate (granular). 
May to August (Plate XVII, fig. 1). 

In rubbly places, in meadow and rocky mountain slopes (up to 
middle mountain zone).— European USSR: Black Sea Region (very 
rarely), Crimea. Caucasus: Ciscaucasia, eastern and southern 
Transcaucasia, Talysh. Soviet Central Asia: Dzhungaria-Tarbagatai, 
mountainous Turkmenia, Syr Darya, Pamiro-Alai, Tien Shan. General 
distribution: Balkans-Asia Minor. Described from Crimea. Type in 
Leningrad. 


9. G. trichophorum Kar. et Kir. in Bull. Soc. Nat. Mosc. XV 
(1842) 372; Ldb. Fl. Ross. II, 418; Fedch. Rast. Turkest. 711. 

Annual. Stem weak, ascending, straight, 15—40 cm tall, repeatedly 
divaricately branched from base, 4-angled almost to top, more or less 
scabrous along ribs due to dense short retrorse setae, often present in 
two rows. Leaves in whorls of 4—8, linear-lanceolate, (5)10-—12(13) 
mm long, (0.5)1—1.5(2) mm broad, gradually acuminate at apex with 
more or less long cusp, gradually narrowed at base, sessile, single- 


310 


293 


veined, with thick antrorse setae along margin and vein beneath, hence 
scabrous; floral leaves in whorls of 4—6, uppermost paired or solitary, 
narrow, (2)3(4) mm long, (0.15)0.25(0.5) mm broad, similar to cauline 
leaves, scabrous. Cymes axillary, numerous in axils of all ramiform 
branches. Peduncles doubly ternately branched, trichoid, long, diffusely 
setulose. Pedicels (4)7—10(16) mm long, glabrous, clavate at top. 
Corolla white-green, small, 1-1.5 mm across, 2-3 mm long, oblong- 
elliptic lobes, with as long slightly fragile cusp at tip. Anthers yellow, 
with filaments shorter than lobes. Style bipartite almost from the base. 
Fruits binate or often 1 mericarp rudimentary, 1.5 mm long, 1 mm 
broad, falcate, barely oblong, glabrous, smooth (fine granulation visible 
only under a binocular microscope) (Plate XVII, fig. 2). 

On stony screes and slopes, rocks, in lower montane belt, 
premontane steppe.—Soviet Central Asia: Balkhash Region, 
Dzhungaria-Tarbagatai, Tien Shan. Endemic. Described from river 
Lepsa (Dzhungarian Ala Tau). Type in Leningrad. 


10. G. transcaucasicum Stapf in Denkschr. Acad. Wissensch. I 
(1885) 52.—G. subreflexum M. Pop. in Tr. prik. bot. gen. and sel. 
XVII, | (1927) 270. (Russian description).—G. anglicum auct. fl. cauc. 
non Huds.—Ic.: M. Pop. I.c. 270, fig. III. 

Annual. Stem filiform, slender, 5-30 cm tall, with thin short 
branches only at base, 4-angled, retrorsely scabrous along ribs. Leaves 
in whorls of 6, upper ones in whorls of 4, at top single and paired, 
oblong, (5)6—7(8) mm long, (0.5)1(2) mm broad, acute, with short 
mucro at apex, almost sessile, retrorsely spinulose along margin and 
vein beneath. Cymes axillary, with 3, rarely with 1 or 2 flowers. 
Peduncles longer than leaves, slender, easily recurved, glabrous, 
sometimes forked (in which case, 6 flowers on 1 peduncle); pedicels 
short, equaling flowers on somewhat longer, recurved in fruit, glabrous. 
Corolla white-yellow, equaling ovary or somewhat smaller, 1.5—2 mm 
across, with 4 ovate lobes with short mucro at tip. Anthers yellow. 
Style bipartite almost from middle. Fruits binate, 1 mm long, 1.5 mm 
broad; mericarp globose, very finely granular-tuberculate, glabrous. 
April to May. 

In thickets of ephemerals, on clayey slopes of mountains, in waste 
places and adyrs, (low foothills bordering the Fergana depression). — 
Caucasus: Dagestan, eastern and southern Transcaucasia, Talysh. Soviet 
Central Asia: Syr Darya, Pamiro-Alai, Tien Shan. Endemic. Described 
from vicinity of Kirovabad (formerly Elizavetpol). Type in Vienna. 

Note. In the herbarium, C.A. von Meyer marked this species for 
description as G. minutiflorum Mey. from Lenkoran. It is distinguished 


31 


— 


294 


from G. anglicum Huds., to which the Caucasian botanists have usually 
referred it, by considerably less branched and often simple stem, long 
peduncles bearing three flowers (and no more), ramose, twice-furcate 
inflorescence and smaller corolla (often smaller than ovary). 

We have not seen the type of this species, but have an adequate 
number of specimens from the classical locality, which completely fit 
the description of this species of Stapf. 


11. G. chloroleucum Fisch. et Mey. in Ind. I sem. Horti Petrop. 
(1835) 27; Linnaea, X, 89; Boiss. FI. or. III, 75; Grossh. Fl. Kavk. IV, 
29. 

Annual. Stems weak, 5-10 cm tall, lodged, highly spreading 
branches, 4-angled, with scattered patent stiff hairs. Leaves in whorls 
of 6-7, oblong-linear or spatulate, 8-10 mm long, 1—1.5 mm broad, 
obtuse or short-acuminate at apex, narrowed toward base, thicker, 
single-veined, patently hispidulus on both surfaces especially along 
margin; floral leaves in whorls of 3 or opposite, smaller than cauline 
leaves. Cymes axillary and terminal, 3—4-flowered. Peduncles almost 
equaling floral leaves. Pedicels 2—3 times as long as flowers, thickended 
in fruit, more or less densely hispidulus. Corolla dirty yellow, 2.5-3 
mm across, with oblong or ovate short-acuminate lobes. Anthers yellow. 
Style bipartite almost from base. Fruits 1 mm long, 2 mm broad, 
mericarps connate at base, binate, rarely with one globose mericarp, 
covered with straight, very short, rigid hairs. June. 

On stony slopes.—Caucasus: southern Ciscaucasia. General 
distribution: Armenia-Kurdistan. Described from neighborhood of Mt. 
Khoi. Type in Leningrad. 

Note. In general appearance it is very similar to G. nigricans 
Boiss., described from Mesopotamia and widespread in Northern Iran, 
from which it is distinguished by the short-acuminate, corolla lobes 
and the rigid, spreading pubescence of the stem. In G. nigricans Boiss. 
the lobes of the corolla have long mucros equaling them in length and 
stems retrorsely scabrous stems. 


Series 2. Rubriflora Pobed.—Flowers purple. 

Note. In Russian floras G. parisiense L. is often reported, which 
belongs to this series. In the Herbarium of the Botanical Institute, 
Academy of Sciences, USSR there is a specimen of this species 
collected by Steven in the Crimea. In the subsequent years, this species 
has not been collected by anyone in the Crimea. It is possible that it 
was introduced and later disappeared. It is also possible that labels 
were mixed up. D.P. Syreistschikov (Fl. Mosk. gub. III (1910) 191) 
reports it as an introduction for the Moscow region, according to 


312 


295 


collections of Maksimovich from Vorobyevy Gory, with two forms: 
trachycarpum Tausch. with pubescent fruits and leiocarpum Tausch. 
with glabrous fruits. 


12. G. decaisnei Boiss. Diagn. ser. I, 6 (1845) 69 and FI or. III, 
77; Trautv. in Tr. SPb. bot. sada, IX, IV, 389; Fedch. Rast. Turkest. 
711; Grossh. Fl. Kavk. II, 30.—G. capillare auct. non Cav.: Decne in 
Ann. Sc. Nat. ser. II, 2 (1834) 269.—Ic.: Post, Fl. Syr. Pal. Sin. 1 
(1932) f. 388.—Exs.: Kotschy, Pl. Pers. bor. aust. No. 66. 

Annual. Stem slender, filiform, 5—25 cm tall, with long, slender 
divaricate branches from base, glabrous, 4-angled. Leaves in whorls 
of 4—6, unequal, linear, very narrow, (9)10—12(15) mm long, 0.25— 
0.5(1) mm broad, single-veined, with revolute margin with or without 
a few white cilia. Cymes axillary, on very slender, glabrous, peduncles 
and pedicels, few-flowered. Peduncles densely-triply furcate or ternate, 
several times as long as leaves. Pedicels 3-6 or more times as long as 
flowers. Corolla purple, almost equaling ovary, 0.5—1 mm across, with 
ovate, short-acuminate lobes. Anthers yellow. Style bipartite from 
middle. Fruits binate, 0.5 mm long, 1 mm broad, glabrous or with 
short scattered white hairs. April to July. 

On limestone rocks, red sandstones, loess mountain slopes, in 
lower montane belt, in shade.— Caucasus: Southern Transcaucasia. 
Soviet Central Asia: Tien Shan, Pamiro-Alai, mountainous Turkmenia. 
General distribution: Balkans-Asia Minor, Armenia-Kurdistan, Iran. 
Described from Southern Iran. Type in Geneva, isotype in Leningrad. 

Note. In the Mediterranean Region this species is replaced by G. 
setaceum Lam., which reaches the eastern limits of its range in Asia 
Minor (Syria, Mesopotamia), but with fruits densely pubescent with 
long erect hairs hooked at the tip. A.A. Grossheim cites G. setaceum 
Lam. for the flora of the Caucasus, apparently only because of an 
erroneous literature record. We did not see a single specimen of G. 
setaceum Lam. from the Caucasus and consider its occurrence in the 
Caucasus doubtful. 


13. G. eldaricum Grossh. in Izv. Azerb. fil. AN SSSR, No. 10, 
(1944) 45; Grossh. Opred. rast. Kavk. 250. 

Annual. Stem slender, filiform, 10—20 cm tall, branched, straight, 
fragile, 4-angled, retrorsely setulose below along ribs, elsewhere 
glabrous, branched, branches erect at tip. Leaves in whorls of 6, middle 
cauline leaves linear , 5-8 mm long, 0.25—0.5 mm broad, upper leaves 
thinner and longer, 8-12 mm long, exceeding inflorescence, sessile, 
with very short mucro at apex, lowest leaves in whorls of 4, obovate, 
attenuated at base into short petiole, acute, all leaves green and 


313 


296 


antrorsely setulose on upper surface, pale green or whitish beneath, 
revolute along margin. Cymes axillary, on very slender glabrous 
peduncles and pedicels, few-flowered. Peduncles doubly-triply furcate 
or ternate, almost equaling leaves. Pedicels 3-6 times as long as 
flowers. Corolla reddish, smaller and narrower than ovary, 0.5 mm 
across, with ovate-oblong, short-acuminate lobes having incurved tip. 
Stamens yellow, shorter than corolla. Style bipartite from middle. Fruits 
binate, 0.5 mm long, 1 mm broad, densely white pubescent. April to 
May (Plate XVII, fig. 5). 

On rubbly-stony slopes.— Caucasus: Eastern Transcaucasia. 
Described from Mt. Eilar-Ougi (Azerbaidzhan). Type in Baku, isotype ~ 
in Leningrad. 


Series 4. Asperae (Moench) DC. Prodr. II (1830) 610, pro 
subsect.—Aspera Moench. Meth. pl. (1794) 641 (pro gen.)— 
Pseudovaillantia Lange in Willk. and Lange, Prodr. fl. Hisp. II (1870) 
309 (pro sect.).—Flowers solitary, subsessile or in small axillary cymes, 
bisexual. Mericarps oblong, straight, densely setaceously pubescent or 
glabrous. Leaves single-veined, in whorls of 4-6. Annuals, small herbs 
with erect stems, scabrous along ribs due to retrorse spinules. 


14. G. verticillatum Danth. in Lam. Encycl. méth. II (1788) 585; 
M.B. FI. taur.-cauc. III, 109; Stev. in Nouv. Mém. Soc. Nat. Mosc. I, 
275; DC. Prodr. IV, 610; Ldb. Fl. Ross. II, 421; Boiss. Fl. or. III, 78; 
Schmalh. FI. II, 11; Grossh. Fl. Kavk. IV, 29.—G. murale M.B. l.c. 
I (1808) 105, non All. and DC.—G. verticillatum Danth. B. leiocarpum 
Rgl. in Bull. Soc. Nat. Mosc. XL, No. 1 (1867) 11.—Sherardia muralis 
Georgi, Beschr. Russ. Reich, IV, 727; Pall. Ind. 105, nom. nud.—Ic.: 
Rchb. Ic. Fl. Germ. XVII, tab. 1201.—Exs.: Herb. Fl. Cauc. No. 245. 

Annual. Stems solitary, 2-5, rarely up to 10, 5-25 cm tall, simple 
or weakly branched, straight or somewhat ascending, 4-angled, scabrous 
along ribs due to short crowded retrorse spinules or in addition, with 
dense, patent hairs along edges. Upper leaves paired, middle leaves in 
whorls of 4 or 6, oblong-linear, (4)5(8) mm long, 0.5-2.5 mm broad, 
with cusp at apex, revolute along margin and scabrous due to dense 
spinules, entire upper surface often spinulose or glabrous (f. 
leiocarpum Rgl.), single-veined, reflexed after anthesis, with very short 
petioles or almost sessile. Flowers in whorls of 4—6 on straight, short, 
thick pedicels; pedicels not exceeding mericarps in length, pubescent 
or glabrous and erect in fruit. Corolla lobes yellow, erect at tip, several 
times as short as ovary, ovate, obtuse, setose outside or only along 
margin or entirely glabrous. Fruits binate, oblong, mericarps arcuate, 
1 mm long, 0.25 mm broad, setosely pubescent or glabrous (f. 
leiocarpum Rgl.). April to September. 


314 


297 


On stony steppes and mountain slopes in foothills.— European 
USSR: Black Sea Region, Crimea. Caucasus: Ciscaucasia, Dagestan, 
eastern and southern Transcaucasia, Talysh. Soviet Central Asia: 
Balkhash Region, mountainous Turkmenia, Syr Darya, Pamiro-Alai, 
Tien Shan. General distribution: Western and eastern Mediterranean, 
Balkans-Asia Minor, Armenia-Kurdistan, Iran. Described from Sicily. 
Type in Paris. . 

Note. The glabrous-fruited form, named B. leiocarpum by Regel, 
is the most widely distributed in Soviet Central Asia, where, sometimes, 
as for example in Turkmenia, it predominates, whereas the form with 
pubescent fruits occurs very rarely. In the Caucasus, the glabrous- 
fruited form is rarer than the pubescent one in Soviet Central Asia. 
Evidently, the form with glabrous fruits does not occur at all in the 
Crimea. 


15. G. apsheronicum Pobed. sp. n. in Addenda XXII, 710. 

Annual. Stems 2-3, 5—18 cm tall, branched at base, straight, or 
somewhat ascending almost terete or barely 4-angled, with weakly 
expressed ribs, glabrous. Leaves in whorls of 6, oblong-linear, 3-6 
mm long, 0.5—1 mm broad, with cusp at apex, scarcely revolute or flat 
along margin and antrorsely spinulose, rarely diffusely spinulose on 
upper surface, single-veined, reflexed after anthesis, with very short 
petioles or almost sessile. Flower axillary in whorls of 3—5, on straight, 
glabrous pedicels; at anthesis pedicels short and thick, elongated and 
recurved in fruit such that mericarps almost press against stem. Corolla 
lobes yellow, at tip straight, 0.5 mm long, several times as short as 
ovary, ovate, obtuse. Stamens 2 times as short as lobes. Style short, 
included in flower. Fruits often with one mericarp, rarely binate, 
mericarps elongated, narrow, arcuate, 1.5—2 mm long, 0.5 mm broad, 
setosely pubescent at tip and with band on one side. Flowering May. 
Fruiting September. 

On coastal sands.— Caucasus: Eastern Transcaucasia. Endemic. 
Described from Apsheron Peninsula. Type in Leningrad. 

Note. Probably, the distribution of this small plant, in nature hard 
to distinguish from G. verticillatum, is more widespread; in any case, 
one can be assured that along the coasts of Caspian Sea it is distributed 
not only in the Apsheron Peninsula, although it has not been collected 
by botanists. 


16. G. nupercreatum M. Pop. in Tr. prikl. bot. gen. i sel. XVII, 
1 (1927) 278.—Ic.: M. Pop. l.c. 270, fig. II. 

Annual. Stem filiform, slender, 14-18 cm long, over entire length 
with few, slender, spreading branches, 4-angled, along ribs smooth or 


317 


298 


hardly scabrous. Leaves lower and upper in whorls of 4, middle in 
whorls of 6, uppermost leaves 1-3, oblong-linear, (3)4(5) mm long, 
0.5—1 mm broad, with short cusp at apex, sessile, short, slender, 
antrorse scattered setae along margin and vein beneath. Cymes axillary, 
1—2-flowered, simple or bifurcate at tip, short, 1.5-2.5 mm long, 
equaling leaves or somewhat shorter, 2—2% times as long as mericarps, 
thin, horizontal, easily arcuately recurved, glabrous. Corolla shorter 
than ovary, whitish-greenish, 0.5—-1 mm across, with ovate, acute lobes, 
without mucro. Anthers yellow. Style bipartite only at top. Fruits black, 
binate, 1 mm long, 0.5 mm broad, mericarps arcuate, narrow, their 
length 2-2% times their breadth, glabrous. May to June (Plate XXI, 
figs’). 

On sandy-pebbly terraces, in ephemeral undergrowth.— Soviet 
Central Asia: Pamiro-Alai. Endemic. Described from Dzham Valley 
(near Samarkand). Type in Tashkent. 

Note. Judging from-some specimens from the vicinity of Stalinabad, 
this species also has a perennial form (collections of Yu.S. Grigoriev 
on sandy-pebbly terrace of Varzob River in the vicinity of Stalinabad, 
18, VI, 1942). 


Section 5. CructaTaE DC. Prodr. IV (1830) 605; Lange in Willk. 
and Lange, Prodr. fl. Hisp. II (1870) 309.—Marschaligalia DC. l.c. 
pro sect.—Cymes axillary, few-flowered. Flowers polygamous, 
unilocular, lower 2—3 in cymes pistillate, rest staminal. Fruits on short, 
recurved pedicels, hidden between leaves. Leaves in whorls of 4, with 
1-3 veins. Perennial, rarely annual herbs with strong, rarely weak 
stems. 


Series 1. Verna Pobed.—Cymes without bracts. 


17. G. vernum Scop. FI. Carn. I (1772) 99; DC. Prodr. IV, 605; 
Ldb. Fl. Ross. II, 417; Boiss. Fl. or. III, 80; Schmalh. Fl. II, 11; 
Grossh. Fl. Kavk. IV, 28; Kryl. Fl. Zap. Sib. X, 2588.—G. polygamum 
Gilib. Fl. lith. I (1785) 3.—G. bauhini Roem. and Schult. Syst. III 
(1818) 218, No. 11.—G. scopolianum Bess. in Schult. Syst. Mant. III 
(1827) 159.—G. glabrum (L.) Roehl ex Steud. Nom. ed. I (1821) 353, 
non Thunb.—G. vernum ssp. caucasicum Ijin in Mat. po istor. fl. i 
rast., vyp. 1 (1941) 260.—Vallantia glabra L. Sp. pl. (1762) 1491; 
Bess. Fl. Galic. I, 117.—Ie.: Scop. l.c. tab. 2; Rchb. Ic. Fl. Germ. 
XVII, tab. 1185; Hegi, Ill. Fl. VI, 1, 226.—Exs.: GRF, No. 175; FI. 
Cechoslov. exs. No. 84; Fl. exs. Reipubl. Boh. Slov. exs. No. 476; FI. 
exs. austro-hung. No. 3749; Doérfler, Herb. norm. No. 5084; Fl. Pol. 
exs. No. 347; Fl. stir. exs. No. 565; Gerb. fl. Ukr. No. 96; Fl. Boh. 


315 


299 


Plate XVII. 
1. Galium tenuissimum M.B., general appearance, part of stem, corolla.—2. G. 
trichophorum Kar. and Kir. corolla—3. G. tianschanicum M. Pop., general 
appearance, ovary, corolla— 4. G. sosnowskyi Mand., general appearance, part 
of stem, leaf—5. G. eldaricum Grossh., general appearance, corolla, fruit.— 
6. G. kaipazii Manden., general appearance, corolla, fruit. 


318 


300 


and Morav. exs. No. 386 [sub. G. cruciata (L.) Scop.]; Fl. Ital. exs. 
No. 1952. 

Perennial. Rootstock slender, creeping, branched. Stems numerous, 
8-50 cm tall, erect ascending or at base, simple, 4-angled, glabrous, 
rarely with patent hairs in lower internodes, with ring of hairs at 
nodes. Leaves in distant whorls of 4, elliptic or oblong-elliptic, obtuse, 
(6)15(20) mm long, (2.5)5(7) mm broad, single-veined, sessile or on 
very short, indistinct petioles, glabrous on upper surface, serrated with 
short antrorse hairs along margin and central vein beneath, reflexed in 
fruit. Cymes axillary, shorter than leaves, 5-flowered, without bracts. 
Peduncles and pedicels glabrous, erect, subsequently recurved. Flowers 
greenish-yellowish, 2 mm across, with elliptic, obtuse or short- 
acuminate lobes, polygamous, lower flowers in each cyme usually 
pistillate, upper staminal fruitless. Style at base with very small discoid 
nectary. Fruits with one mericarp, globose-reniform, at base somewhat 
deflexed, 1.5-2.5 mm in broad part, glabrous, smooth. April to August. 

In open pine, birch and beech forests, on stony screes, in groves, 
in alpine and subalpine meadows.— European USSR: Upper Dnieper, 
Middle Dnieper, Volga-Don, Upper Dniester, Black Sea Region. 
Caucasus: Ciscaucasia, Dagestan, eastern Transcaucasia. Western 
Siberia: Altai. General distribution: Central Europe, western and eastern 
Mediterranean, Balkans-Asia Minor, North America (introduced). 
Described from Austria. Type in London. 

Note. In the Caucasus this species is more often found in the 
subalpine and alpine zones of the Greater Caucasian Range and in 
Dagestan. It differs from the Western European species in having a 
small stature, small leaves and short internodes, so that whorls of 
leaves are strorgly telescoped. 

M.M. Iljin assigned the alpine form, in our opinion, only a 
Caucasian ecological form, to the rank of subspecies [Mat. po istor. 
fl. i rast., vyp. 1 (1941) 260]. In the Caucasus one often finds hybrids 
between G. vernum Scop. and G. cruciata (L.) Scop. with varying 
combinations of characters; in eastern Transcaucasia, there is typical 
G. vernum Scop. 

We did not find specimens of this species from the Altai, but L.P. 
Sergievskii has confirmed that in the herbarium of Tomsk University 
there are preserved collections of Krylov from a single locality of 
conifer forest in the “saddle” between the upper reaches of the Kytma 
River and the peak of Chernaya Uba. Evidently, this is a very rare, 
relict locality of G. vernum in the Altai, since collections of it have 
not be made again in the Altai. 


18. G. schischkinii Pobed. sp. n in Addenda XXII, 711. 


301 


Perennial. Root slender, reddish brown. Stem ascending, straight, 
5-6 cm long, weakly branched, 4-angled, with dense, retrorse spinules 
on lower internodes along ribs, moreover with long, white soft hairs. 
Leaves in whorls of 4, elliptic or lanceolate, 5-7 mm long, 2.5-4 mm 
broad, subobtuse at apex, gradually narrowed toward base, sessile, 
with 3 indistinct veins, with long, white, antrorse cilia along margin, 
with still longer, white hairs along middle vein beneath. Cymes axillary, 
4 in each of upper whorls, 3-flowered and several rudimentary 
underdeveloped cymes in each whorl, cymes in axils of lower whorls 
paired, single-flowered. Flowers all bisexual. Peduncles 2—3 times as 
short as leaves, covered by very long white hairs. Pedicels equaling 
flowers, with clusters of long hairs, exceeding flower, recurved in 
fruit. Corolla 1-1.5 mm across, evidently greenish, with elliptic, obtuse 
lobes. Anthers yellow. Style bipartite almost from base. Fruits 
(immature) binate, glabrous, smooth. 

Banks of saline lakes.— Caucasus: Eastern Transcaucasia. Endemic. 
Described from environs of Tbilisi (Kuky salt lakes). Type in 
Leningrad. 

Note. This species is distinct in the series because of its dense 
retrorse spinules on the stem. In this character, it very closely resembles 
G. vernum, but in its pubescence on the lower internodes and long 
white hairs on the leaves, it resembles G. cruciata. It is distinguished 
from both species in having extremely short peduncles (2-3 times 
shorter than leaves); pedicels covered with long white hairs, and also 
a reduction of the cymes in each whorl to four in the upper internodes 
and two in the lower internodes. 


19. G. krylovii Iljin in Animadvers. syst. ex Herb. Univ. Tomsk. 
No. 5 (1935) 1; Kryl. Fl. Zap. Sib. X, 2588. 
Perennial. Rootstock slender, branched. Stems numerous, 10-40 
cm tall, simple or rarely with isolated branches closer to base, 4- 
angled, pubescent with long erect hairs, glabrous, occasionally in lower 
part. Leaves in whorls of 4, more or less distant, oblong-elliptic, 
(8)16(25) mm long, (4)7—8(9) mm broad, obtuse, sessile, with 3 
indistinct veins, with long rigid hairs beneath or only along middle 
vein and margin. Cymes 5-flowered, axillary, shorter than leaves, 
without bracts, peduncles and pedicels glabrous, lower flowers 
pistillate, upper staminal. Corolla 3 mm across, its lobes oblong-elliptic, 
subobtuse. Style at base with discoid nectary. Fruits with one mericarp, 
globose or reniform, 2 mm broad, glabrous. May to August (Plate 
XVI, fig. 4). 
On mountain slopes: in forest meadows, scrubs, and pine, fir, 
319 spruce and larch forests (in black taiga).— Western Siberia: Ob’, Altai. 


302 


Eastern Siberia: Angara-Sayan. General distribution: India- 
Himalayas, Mongolia. Described from Minusinsk district. Type in 
Leningrad. 


20. G. elbrussicum Pobed. sp. n. in Addenda XXII, 711. 

Perennial. Root filiform, long, branched, dark brown. Stems one 
or several from root, weak, lodged or ascending, 5—7 cm tall, with 
divaricate, almost horizontal, long branches, 4-angled, lower internodes 
glabrous, elsewhere densely pubescent with patent hairs. Leaves in 

- whorls of 4 (whorls crowded on stem, internodes shorter than leaves), 

elliptic, 4-5 mm long, 2-3 mm broad, obtuse or almost rounded at 
apex, at base barely narrowed into very short petiole or almost sessile, 
thicker, with one distinct vein (reticulate branching of vein visible 
only under high-magnification binoculars), ciliate along margin and 
with scattered, appressed hairs on upper surface. Cymes in axils of 3- 
6 upper whorls .of leaves, few-flowered. Peduncles doubly ternately 
and furcately branched, without bracts, with dense, patent hairs together 
with pedicels recurved in fruit. Flowers polygamous, lower ones in 
cyme fruit-bearing, upper ones staminal without fruit. Corolla 2 mm 
across, with oblong, obtuse lobes with dark brown middle and marginal 
veins. Anthers yellow, on long filaments, almost equaling lobes. Style 
bipartite. Fruit globose, large, 3 mm in diameter, with one mericarp, 
rugose, glabrous, young fruit with distinct discoid nectary in the form 
of an annulus. September. 

On screes in alpine zone.— Caucasus: Western Transcaucasia. 
Endemic. Described from Mt. Elbrus. Type in Leningrad. 


Series 2. Cruciata Pobed—Cymes with bracts. 


21. G. cruciata (L.) Scop. Fl. Carn. ed. I, 1 (1772) 100; DC. 
Prodr. IV, 606 ; Ldb. FI. Ross. II, 416; Boiss. Fl. or. III, 79; Grossh. 
Fl. Kavk. IV, 28.—G. vernum B. cruciata Schmalh. FI. II (1897) 12. 
—G. chersonense auct. non. Willd.—Valantia cruciata L. Sp. pl. 
(1753) 1052; Georgi, Beschr. Russ. Reich. III, 4, 1366; Steven in 
Mém. Soc. Nat. Mosc. I, 275.—V. hirsuta Gilib. Exerc. phyt. I (1792) 
25.— V. hispida Georgi, |.c. (1800) (tantum pl. lithuanica).—Ic.: Rchb. 
Ic. Fl. Germ. XVII, tab. 1185; Fedch. and Fler. Fl. fig. 898; Hegi, Ill. 
Fl. VI, 1, tab. 248.—Exs.: Fl. exs. austro-hung. No. 3745; FI. Siles. 
exs. No. 381; Fl. Gal. and Germ. exs. No. 2475; Pl. exs. Anat. or. No. 
1049; Fl. Stir. exs. No. 1256; Dérfler, Herb. norm. No. 5083 [sub G. 
chersonense (Willd.) Roem. and Schult.]. 

Perennial. Rootstock long, slender, creeping, branched. Stems 

320 numerous, 20-60 cm tall, straight or ascending at base, simple, 


303 


4-angled, densely or diffusely pubescent with long erect hairs, rarely 
glabrous (var. chersonense Roem. et Schult. pro sp.). Leaves 4 in 
distant whorls, oblong-elliptic or elliptic, obtuse or subacute, (8)15(22) 
mm long, (2)5(8) mm broad, sessile, 3-veined, glabrous or diffusely 
pubescent on upper surface, beneath and along margin densely 
pubescent or only ciliate, and with pubescence along middle vein 
(var. chersonense), reflexed and appressed to stem in fruit. Cymes 
axillary, shorter than leaves, 5—6-flowered. Peduncles and pedicels 
pubescent with long patent or appressed hairs; recurved in fruit. Flowers 
in branched cymes, with 2 bracts resembling leaves, but considerably 
smaller, yellow, 2—3 mm across, with ovate acute lobes, one-two lower 
flowers bisexual, fruit-bearing, remaining staminal, without fruit. Style 
at base with discoid nectary. Fruits with one mericarp (rarely with 
underdeveloped second mericarp), globose, smooth, glabrous, 1—1.5 
mm in diameter. May to July. 

On rocks, in subalpine meadows, forest clearings and forest 
glades.— European USSR: Middle Dnieper, Volga-Don, Bessarabia; 
Black Sea Region; Crimea. Caucasus: All regions. General distribution: 
Central Europe, western and eastern Mediterranean, Balkans-Asia 
Minor, Armenia-Kurdistan, Iran-Afghanistan, North America 
(introduced). Described from Germany. Type in London. 

Note. A hybrid between G. cruciata (L.) Scop. and G. 
pedemontanum (Bell.) All. is found in the Caucasus; in habit it 
resembles the former, but in the characteristic pubescence of the stem, 
being retrorse spinules, it resembles the latter species. 


22. G. tauricum (Willd.) Roem. et Schult. Syst. veg. III (1818) 
250; DC. Prodr. IV, 605; Ldb. Fl. Ross. II, 416, p.p.—G. coronatum 
auct. non Sibth. and Sm.: Boiss. Fl. or. III (1875) 79; p.p.; Schmalh. 
Fl. II, 12, p.p.—G. coronatum Sibth. et Sm. var. lasiocarpum Boiss. 
in Sched.—G. cruciata var. taurica Trautv. in Tr. SPb. bot. sada, IV 
(1876) 145.—G. humifusum Grossh. in Fl. Kavk. IV (1934) 28, p.p.— 
Valantia taurica Willd. Sp. pl. TV (1806) 951; M.B. FI. taur.-cauc. II, 
436, p.p. (tantum pl. taur.).— V. chersonense Willd. Sp. pl. IV (1806) 
851.—Exs.: GRF, No. 669, p.p.; Dérfler, Herb. norm. No. 4532. 

Perennial. Rootstock vertical, slender, branched, woody. Stems 
numerous, 7-30 cm tall, ascending, branched at base, woody, densely 
pubescent with long erect hairs. Leaves in whorls of 4, obovate or 
elliptic, obtuse or short-acuminate, narrowed at base into short petiole 
or sessile, lower ones very tiny, crowded, middle leaves (5)7(10) mm 
long, (2.5)4(6) mm broad, more or less distant, upper ones often 
suborbicular, crowded, on vegetative shoots almost linear, narrow, 8— 
14 mm long, 1.5-3 mm broad, all leaves densely pubescent on both 


— 


304 


surfaces or with cilia only along margin and middle vein beneath, 
rarely entirely glabrous, with one vein, entire. Cymes axillary, 5-6 
flowered. Peduncles longer than leaves, together with pedicels densely 
pubescent with long patent hairs disappearing after anthesis, with 
elliptic, long-ciliate bracts. Flowers polygamous, usually 2 lower- 
pistillate remaining staminal. Corolla yellow, 3 mm across, with ovate 
or elliptic, acute or subobtuse lobes. Style with discoid nectary at 
base. Fruits with one mericarp, globose, 3—3.5 mm in diameter, densely 
covered with short hairs and sparse, long hairs or entirely glabrous. 
April to July (Plate XVI, fig. 2). 

On rocks, steppe slopes, in scrubs.— European USSR: Crimea. 
Caucasus: Ciscaucasia, western Transcaucasia. General distribution: 
Balkans-Asia Minor (regions of Asia Minor, Cilicia, Lydia, 
Mesopotamia, Syria, Anatolia), Armenia-Kurdistan, Iran. Described 
from Crimea. Type in London. 

Note. It should be observed that strongly pubescent plants with 
densely, short-pubescent fruits occur only in the Crimea, whereas an 
entirely glabrous race is found only in the Caucasus, which means that 
G. tauricum and G. coronatum have strictly isolated geographical 
ranges. The form with long, ciliate leaves, bracts and peduncles, but 
with glabrous fruit is widespread not only in the Crimea, but also in 
the northern Caucasus, where forms with pubescent fruits are not seen, 
which forces us to refer to this glabrous-fruited form as most likely a 
hybrid species, having developed in the past when both the parent 
species G. tauricum and G. coronatum grew in the same territory. We 
cannot recognize this hybrid as a separate species, as was done by 
Willdenow, giving it the name Valantia chersonensis (G. chersonense), 
since its characters are not stable. 


23. G. sosnowskyi Manden. in Zam. po sist. i geogr. rast. Tbilissk. 
bot. inst. 13 (1947) 114; Grossh. Opred. rast. Kavk. 249.—G. 
coronatum f. isophyllum Boiss. Fl. or III (1875) 80. 

Perennial. Plant densely patently pubescent. Stems numerous, 
slender, procumbent, 5—7 cm tall. Leaves in whorls of 4, ovate or 
ovate-oblong, very small, 4-6 mm long, 2-3 mm broad, single-veined, 
densely distributed on stem, covering internodes. Cymes axillary, 
almost equaling leaves, with 2-3 ovate or ovate-oblong floral leaves 
at base. Flowers polygamous, 2-3 lower pistillate, remaining staminal, 
fruitless. Corolla golden yellow, 2.5 mm across, with ovate-oblong, 
short-acuminate lobes. Peduncle and pedicels arcuately recurved after 
anthesis. Mericarp single, glabrous, 3 mm in diameter. July (Plate 
XVII, fig. 4). 

On volcanic screes.— Caucasus: southern Transcaucasia. Endemic? 
Described from Armenia. Type in Tbilisi. 


322 


305 


Note. The isotype of the species most closely approaches G. 
tauricum (Willd.) Roem. et Schult. and not any other species with 
which the author has compared it while describing it. The fundamental 
distinguishing characters between G. sosnowskyi and G. tauricum are: 
ovate shape of leaves, length of pedicels, equaling tiny leaves, and 
glabrous fruits, which are not typical of G. tauricum and are found 
only in hybrids between G. tauricum and G. coronatum. 


24. G. braunii Zelen. in Mat. dlya Flory Kryma (1906) 284.—G. 
coronatum var. braunii (Zelen.) Fedtsch. and Fler. Fl. (1910) 912. 

Perennial. Stems more or less woody at base, numerous, decument 
or ascending, simple, glabrous (var. glaberrimum Zelen.) or covered 
with long, soft hairs (var. hirsutissimum Zelen.), with shortened 
internodes at base. Leaves in whorls of 4, very small, obovate, obtuse, 
2-3 mm long, 1-1.5 mm broad, single-veined, glabrous (var. 
glaberrimum Zelen.), ciliate along margin or hairy on both surfaces 
(var. hirsutissimum Zelen.), lower ones finer. Cymes axillary, 
approximate, much shorter than leaves, with somewhat larger flowers, 
recurved, with bracts, branched, glabrous. Corolla yellow, with ovate, 
acute lobes. Fruit unknown. April. 

In steppe regions of Crimea.— European USSR: Crimea. Described 
from vicinity of Simferopol. Type in Nikitsky Garden. 

Note. Without comparing specimens of this species with G. 
tauricum and G. coronatum, we cannot assert with conviction its 
independent status, more so because G. tauricum M.B. also occurs 
near Simferopol. 


25. G. coronatum Sibth. et Sm. Fl. Graec. I (1806) 90; DC. 
Prodr. IV, 605; Ldb. Fl. Ross. II, 416; Boiss. Fl. or. III, 79; Schmalh. 
Fl. I, 12.—Valantia humifusa Willd. Sp. pl. II (1805) 949; M.B. FI. 
taur.-cauc. II, 436 and III, 640.—G. coronatum f. glaberrimum DC. 
Prodr. IV (1830) 605.—G. coronatum y. stenophyllum Boiss. FI. or. 
III (1875) 80.—G. cruciata var. humifusa Trautv. in Tr. SPb. bot. 
sada, II (1873) 542.—G. humifusum (Willd.) Stapf. in Denkschr. Akad. 
Wissensch. Wien, 50, I (1885) 35, non M.B.; Grossh. Fl. Kavk. IV, 
28.—G. persicum DC. l.c. 605.—Ic.: Sibtorp and Sm. lL.c. II, tab. 
125.—Exs.: Pl. exs. Anatol. or. No. 931. 

Perennial. Rootstock vertical, branched, woody. Stems numerous, 
4—35 cm tall, straight or ascending, simple, somewhat branched and 
woody only at base, glabrous. Leaves in whorls of 4, obovate, or 
obovate-orbicular, (5)10-12(20) mm long, (2)6—-7(10) mm broad, 
obtuse or rounded at apex, sessile thicker, single-veined, rarely 3- 
veined, glabrous, more rarely with cilia along margin, lower leaves 


323 


306 


very small, aggregated, cauline and upper leaves larger, more or less 
evenly distributed, leaves of vegetative shoots narrower, elliptic to 
linear, 11-20 mm long, 2-3 mm broad. Cymes axillary 5—6-flowered. 
Peduncles and pedicels glabrous, equaling or longer than leaves, 
recurved in fruit. Flowers yellow, polygamous, 2 lower flowers in 
cyme usually bisexual, remaining staminal, 3-3.5 mm across, their 
lobes lanceolate or elliptic. Fruit with single mericarp, globose, 
glabrous, 3 mm in diameter. April to August. 

On dry, rubbly slopes and in scrubs.—Caucasus: Ciscaucasia, 
Dagestan, western, eastern and southern Transcaucasia. General 
distribution: Balkans-Asia Minor, Armenia-Kurdistan, Iran. Described 
from Cappadocia. Type in London. 

Note. The lone report on the occurrence of this species in the 
Crimea (Baidar Pass), unconfirmed since, causes us to suggest that 
labels could have been confused, because it appears to us very doubtful 
that this species was growing in Crimea. 


26. G. kopetdaghense Pobed. sp. n. in Addenda XXII, 711.—G. 
coronatum auct. fl. As. med.—Valantia taurica auct. fl. As. med. 

Perennial. Root vertical, thick, branched, woody, forming caudex. 
Stems numerous, woody at base, 12-30 cm tall, simple, branching 
only at base, densely, softly pubescent or glabrous, same for entire 
plant (var. glabrescens m.). Leaves in whorls of 4, more or less evenly 
distributed, lower leaves smaller, aggregated, broadly elliptic, obovate 
or almost orbicular, (8)13(21) mm long, 8-14 mm broad, obtuse, 
sessile, thicker, 1-3 veined, uniformly pubescent with short, antrorsely 
appressed hairs, leaves of vegetative shoots narrowly elliptic, 7-25 
mm long, 2.6 mm broad, obtuse. Cymes axillary, longer than leaves 
after anthesis. Peduncles and pedicels short pubescent with patent 
hairs. Bracts smaller than leaves, do not differ from leaves in shape 
and pubescence. Flowers yellow, polygamous, 3—4 mm across, their 
lobes elliptic-oblong, obtuse. Fruits globose, 3—3.5 mm in diameter, 
with scattered, more or less long hairs. April to June (Plate XVI, fig. 3). 

On steppe slopes of mountains.— Soviet Central Asia: mountainous 
Turkmenia. Described from Kopet Dag. General distribution: Iran- 
Afghanistan (north)? Type in Leningrad. 

Note. This species is extremely close to G. tauricum (Willd.) Roem. 
et Schult.; however, it is well-distinguished by its evenly distributed 
(and not aggregated at top of the stem) leaf whorls, soft, shorter stem 
pubescence (and not rigid, long, patent), short, antrorsely appressed 
pubescence (and not long, spreading) of the leaves and peduncles, 
fruit covered by scattered, more or less long (and not thick, short) 
hairs, and considerably thicker root. If one takes into consideration 


324 


307 


the wide separation of the ranges of these comparable species (Crimea- 
Kopet Dag), then isolating them as well-expressed geographical race 
is indisputable. 


27. G. sevanense Pobed. sp. n. in Addenda XXII, 712. 

Perennial. Root filiform, long, reddish brown. Stems several with 
root, 5-8 cm tall, branched from base, 4-angled, more than lower half 
glabrous, upward pubescent with patent hairs. Leaves at bottom very 
small, scalelike in entire part of glabrous stem (evidently, filled with 
loose substratum), upper green, in whorls of 4, almost orbicular or 
orbicular-elliptic, 3-5 mm long, 2.5-3 mm broad, obtuse or rounded 
at apex, thicker, sessile, single-veined, more or less pubescent along 
margin and over entire surface, whorls with green leaves closely packed, 
with lamellar-spacings. Cymes axillary, 5-6 in all green whorls, with 
broadly ovate bracts 1 mm long and 0.5 mm broad, simple, 3-flowered. 
Peduncles equaling leaves or somewhat longer, together with flowers 
glabrous or patently pubescent. Pedicels equaling flowers or shorter. 
Flowers polygamous, lower usually pistillate, upper staminal. Corolla 
2.5—3 mm across, with oblong, obtuse, somewhat unequal (2 decussate, 
slightly narrower than other 2) lobes. Anthers yellow, large; filament 
long, almost equaling lobes. Style bipartite. Mature fruits unknown. 
June. 

On mountain slopes.— Caucasus: Eastern Transcaucasia. Described 
from Lake Sevan (Akhdag Range). Type in Leningrad. 

Note. From G. coronatum Sibth. et Sm. it differs in habit and in 
having very small, scalelike leaves over the greater part of the stem, 
leaf whorls aggregated on the upper part of the stem, and small, thick, 
coriaceous green leaves; the cymes of G. sevanense have three, not 5— 
6 flowers, as in G. coronatum. 


28. G. cordatum Roem. et Schult. Syst. III (1818) 259; DC. Prodr. 
IV, 606.—G. articulatum (L.) Roem. et Schult. I.c. 250; C.A.M. 
Verzeichn. 55, Hohenack. Enum. Elisabethpol. 216; Ldb. Fl. Ross. II, 
417; Boiss. Fl. or. III, 81; Schmalh. Fl. II, 12; Grossh. Fl. Kavk. IV, 
27.—G. pygmaeum DC. Prodr. IV (1830) 606.—Valantia articulata 
L. Sp. Pl. (1753) 1052.—V. muralis M.B. Fl. taur-Cauc. II (1808) 
435, non L.—V. pusilla Stev. in Nouv. Mém. Soc. Nat. Mosc. I (1829) 
274, id. est Mém. Soc. Nat.. Mosc. VI (1829) 274.—Ic.: Pflanzenfam. 
IV, 4, 150.—Exs.: Fl. Palaest. exs. No. 289. 

Annual. Stems one—many, simple or branched, straight, 3-15 cm 
tall, glabrous. Leaves in closely-packed whorls of 4, broadly cordate, 
obtuse, (4.5)8(15) mm long, (2.5)10(12) mm broad, in broader part 
reflexed even at time of flowering, sessile, single-veined, glabrous 
with pubescence only along lower margin and base of middle vein on 


308 


upper surface. Cymes 4—6 flowered, on peduncles considerably shorter 
than leaves, with very tiny papilla, curved under leaves when in fruit. 
Flowers yellow, 1.5—2 mm across, with ovate acute corolla lobes. Fruits 
with single mericarp, reniform, 2 mm in diameter or rarely binate, 3 
mm broad, covered with extremely tiny papillae and often with white, 
linear spots. March to June. 

On dry stony mountain slopes, stony screes.— Caucasus: Eastern 
and southern Transcaucasia, Talysh. General distribution: Eastern 
Mediterranean, Balkans-Asia Minor, Iran. Described from Egypt. Type 
in London. 

Note. In literature there is a report of the occurrence of this species 
in the Crimea; however, we have not seen an authentic specimen of G. 
articulatum from the Crimea. Probably, reported erroneously. 


29. G. pedemontanum (Bell.) All. Auct. Fl. Pedem. (1789) 2; 
DC. Prodr. IV, 605; Ldb. Fl. Ross. II, 417; Boiss. Fl. or. III, 80; 
Schmalh. Fl. II, 11; Grossh. Fl. Kavk. IV, 28.—G. chloranthum Brot. 
Fl. Lusit. I (1804) 149.—G. retrorsum DC. 1.c. 605.—Valantia 
pedemontana Bellardi, App. (1788) 46; Willd. Sp. pl. IV, 949; M.B. 
Fl. taur.-cauc. II, 436.—Iec.: Rchb. Ic. Fl. Germ. XVII, tab. 1185; 
Hegi, Ill. Fl. VI, 1, 226.— Exs.: Herb. Fl. Cauc. No. 244; Fl. Boh. and 
Morav. exs. No. 684; Fl. Ital. exs. No. 663; Fl. Gal. and Germ. exs. 
No. 3399; Fl. Cechoslov. exs. No. 83. 

Annual. Stem single, rarely 3-5, straight, simple, slender, 5-50 
cm tall, retrorsely spinulose along ribs, sometimes mixed with thin, 
long hairs, rarely only the latter. Leaves in widely scattered whorls of 
4, elliptic, tiny, (2)5(11) mm long, (0.75)2(5) mm broad, obtuse, sessile, 
single-veined, more or less densely pubescent with long appressed 
hairs on both surfaces, subsequently reflexed. Cymes 2-3 flowered, 
considerably shorter than leaves. Peduncles and pedicels recurved and 
hidden under leaves in fruit, pubescent with long white hairs. Flowers 
yellow, 1-1.5 mm across, polygamous. Corolla lobes broadly elliptic, 
obtuse. Fruits with single mericarp, occasionally binate (then 2 mm 
broad), globose, 1 mm in diameter, smooth, glabrous. April to July. 

In sandy places along banks of lakes, subalpine meadows, glades 
small meadows, on steppe slopes of mountains, in scrub.— European 
USSR: Bessarabia, Black Sea Region, Crimea. Caucasus: Ciscaucasia, 
Dagestan, western and eastern Transcaucasia, Talysh. Soviet Central 
Asia: Tien Shan (Karatau), Pamiro-Alai. General distribution: Central 
Europe, western and eastern Mediterranean, Balkans-Asia Minor, 
Armenia-Kurdistan, Iran. Described from Italy. Type in Turin. 


326 


309 


30. G. pseudopolycarpon Somm. et Lév. in Bull. Soc. Bot. Ital. 
(1894) 30; Tr. SPb. bot. sada, XVI, 211; Grossh. Opred. rast. Kavk. 
249.—Ic.: in Tr. SPb. bot. sada, tabl. XXII. 

Perennial. Stems filiform, 5.5-6 cm tall, at base bitripartite, 
divaricately branched, fragile, 4-angled, glabrous, with 2.5 cm long 
internodes. Leaves in whorls of 4, unequal, obovate-lanceolate, 8 mm 
long, 4 m broad, obtuse, single-veined, thicker, at base extended into 
short petiole, reflexed, narrowly scarious along. margin and here 
glabrous or sparsely ciliate; cauline leaves larger and broader. Cymes 
terminal and in axils of upper leaves, with 3 floral leaves, simple, 3- 
flowered. Pedicels recurved in mature fruits, with paired bracts at 
base, glabrous, sturdy, all flowers bisexual. Fruits large, 3 mm long, 
2 mm broad, binate or with one mericarp, glabrous. September. 

In alpine mountain belt.—Caucasus: Western Transcaucasia. 
Endemic. Described from Mt. Elbrus. Type in Florence. 

Note. We never saw specimens of this species. I.P. Mandenova 
[Zam. po sist. 1 geogr. rast. Tbilissk. bot. inst. 17 (1953) 62], on the 
basis of the diagnosis and illustrations, referred this species to Asperula 
gracilis (described from Elbrus), with which we are unable to agree. 
According to the structure of the inflorescence, this species is close to 
the genus Galium and not to Asperula, but it is impossible to say this 
with confidence, because we do not know whether the flowers of this 
species possess a tube. Judging from the illustration and diagnosis, 
this species differs from Asperula gracilis C.A.M., in having downward 
recurved fruit stalks in the mature fruits and reflexed cauline leaves. 
In the Herbarium of Botanical Institute of the Academy of Sciences, 
USSR, there is another specimen close to G. pseudopolycarpon from 


Elbrus, misidentified by its collector, Kharkevich, as G. humifusum 


(Willd.) Stapf. 


Section 6. CyMOGALIAE Pobed. sect. n. in Addenda XXII, 712.— 
Trichogalia DC. Prodr. IV (1830) 599, p.p.; pro subsect.—Cymes 
furcate, terminal. Flowers bisexual. Leaves in whorls of 2-4, 1-3 
veined. Fruits pubescent with long, dense, setaceous hooked hairs. 
Perennial herbs with slender, weak, glabrous or somewhat pubescent 
stems. 


Series 1. Paradoxa Pobed.— Leaves in whorls of 4, lowest of lower 
leaves paired, single-veined. 


31. G. paradoxum Maxim. in Bull. Acad. Sc. Pétersb. XIX (1814) 
281; Kom. in Tr. SPb. bot sada, XXV, 495; Kom. and Alis. Opred. 
rast. Dalnevost. kr. II, 948.—G. syreitschikowii Lipsch. in Fedde, 


329 


310 


Repert. 26 (1929) 286. tab. XCVII.—Ic.: Miyoschi and Makino, Pock. 
Atlas Alp. pl. Jap. II (1907), tab. 57, f. 328; Martynenko in DAN 
SSSR, 31, 9 (1941) 959-961. 

Perennial. Rootstock creeping, rooting at nodes. Stems 10-22 cm 
tall, straight, slender, smooth. Leaves in 3 lower nodes paired, orbicular, 
emarginate at apex, pubescent on both surfaces or only upper and 
beneath along veins. with scattered, short appressed hairs, ciliolate 
along margin, with one distinctly noticeable vein, with petioles equaling 
half the length of lamina or shorter, with 2 small lanceolate stipules, 
situated decussately with leaves; higher cauline leaves longer, in whorls 
of 4, 2 of them considerably larger than other 2, ovate, very short- 
acuminate, with pubescence similar to lower leaves. Cymes simple, 3- 
flowered, in threes, terminal stems and branches. Peduncles more or 
less long, with 2 small lanceolate bracts. Pedicels short. Corolla 2.5— 
3 mm across, white, with oblong-elliptic, obtuse lobes. Fruits with 
dense, long hooked, shining setae. June to August. 

In shady places of coniferous and mixed forests.— European USSR: 
Volga-Kama. Western Siberia: Altai. Eastern Siberia: Angara-Sayan 
(Lake Baikal). Soviet Far East: Ussuri. General distribution: India- 
Himalayas, Japan, China. Described from southeastern Manchuria. Type 
in Leningrad. 

Note. The Japanese plants of this species, as already observed 
[Franch. and Savat. Enum. pl. II (1879) 392], differ somewhat from 
the continental plants as follows: lower leaves whorled except 
sometimes at the lowest nodes a whorl is not formed; all leaves 3- 
veined. However, in Japanese plants the three veins are as poorly 
expressed as in continental plants; hence, there are no essential 
differences 

The species is most widespread in the Soviet Far East, China, 
Korea and Japan. Moreover, it is known in disjunct regions apart from 
the indicated geographical range: in the Himalayas, Altai and Urals, 
where it is rare and is found as a relict. 


Series 2. Rotundifolia Pobed.— All leaves in whorls of 4, equal, 
with 3 distinctly prominent veins. 


32. G. scabrum L. Sp. pl. (1753) 108.—G. rotundifolium L. Sp. 
pl. (1753) 108; DC. Prodr. IV, 599; Ldb. Fl. Ross. II, 412; Boiss. FI. 
or. III, 49; Schmalh. Fl. II, 17; Grossh. Fl. Kavk. IV, 30.—Ic.: Hegi, 
Ill. Fl. VI, 1, tab. 248; Bonnier, Fl. tab. 255.—Exs.: Fl. Boh. and 
Morav. exs. No. 194. 

Perennial. Stems numerous, decumbent or ascending, 10-40 cm 
tall, straight, simple, rarely somewhat branched, slender, 4-angled, 


311 


Plate XVIII. 


327 1. Galium karakulense Pobed., upper part of plant, corolla, section of stem.— 
2. G. baicalense Pobed., general appearance, corolla.—3. G. ruprechtii Pobed., 
part of plant, corolla, fruit—4. G. saturejifolia Trev., inflorescence, corolla, 

fruit. 


330 


S12 


smooth, glabrous or with few scattered hairs. Leaves in whorls of 4, 
broadly elliptic, broadly ovate or almost orbicular, (9)15(21) mm long, 
(5)9(10) mm broad, obtuse or often short-acuminate at apex, 3-veined, 
along margin fimbriate with setae, glabrous or diffusely setulosely 
pubescent on both surfaces, pale green, with short petioles. Cymes 
furcately branched or simple, 6-, rarely 3- flowered, in threes, terminal 
on stem, rarely also in axils of upper leaves. Peduncles and pedicels 
straight, glabrous, long, many times as long as flowers. Corolla 3—3.5 
mm across, white or greenish (dry-brownish), with ovate acute lobes. 
Fruits with dense, long hooked setae. June to August (September). 

In coniferous forests (principally fir), less often in beech forests.— 
European USSR: Upper Dniester, Caucasus: Western and eastern 
Transcaucasia, Talysh. General distribution: Scandinavia (southern), 
Central Europe, western and eastern Mediterranean, Balkans-Asia 
Minor, Armenia-Kurdistan. Described from Southern Europe. Type in 
London. 

Note. Ruprecht [Rupr. Fl. ingrica (1860)] indicates that this species 
has been erroneously reported in the literature for the flora of the 
Leningrad Region. 


33. G. kamtschaticum Steller ex Schult. Mant. III (1827) 186; 
Kom. and Alis. Opred. rast. Dalnevost. kr. II, 948; Kom. Fl. Kamch. 
III, 100.—G. obovatum Ldb. FI. Ross. II (1844) 412; non Kunth.— 
Ic.: Britt. and Br. Ill. Fl. N. St. and Canada, III (1898) 222; ed. 2, III 
(1913) 261; Miyoschi and Makino, Pock. Atlas Alp. pl. Jap. II, tab. 
STS 253 

Perennial. Rootstock filiform, creeping, with rooting at nodes, 
semi-subterranean shoots, and scale-like, tiny, opposite leaves, 
blackening on drying. Stem very slender, simple, smooth. Leaves in 
whorls of 4, broadly obovate or orbicular, (14)15—20(25) mm long, 
and (11)12—15(17) mm broad, rounded at apex, mucronate, thin, sessile, 
or with very short petioles, 3-veined, with scattered, short, appressed 
hairs on upper surface and beneath along veins or on both surfaces, 
short ciliate along margin, dark green. Cymes simple or furcately 
branched, 3—6-flowered, in threes, terminal on stems, rarely also in 
axils of upper leaves, on long peduncles, with 2 small, lanceolate 
bracts at base of pedicels. Corolla 3-4 mm across, greenish white 
(brown when dry), with ovate-lanceolate, acute lobes. Fruits with thick, 
long hooked setae. July to September. 

In moss-covered coniferous forests and especially in alder groves 
and birch forests.—Soviet Far East: Kamchatka, Sakhalin (in Kuril 
Islands), Uda, River area, Ussuri. General distribution: Japan, Bering 


33 


— 


318 


Strait, North America. Described from Kamchatka. Type evidently not 
preserved. 

Note. Ledebour did not distinguish this species from the South 
American G. obovatum Kunth [ex H.B. Nov. gen. Am. 3 (1818) 263, 
tab. 277], although the latter differs from G. kamtschaticum by the 
elliptical shape of the leaves and stronger pubescence all over the 
plant, particularly on the stem. All subsequent reports of G. obovatum 
from the eastern part of the Asian Continent and the adjoining islands 
are erroneous. ; 


Section 7. APARINOIDES Jord. Obs. III (1846) 168, p.p.—Leiogalia 
DC. Prodr. IV (1830) 593, p.p.; pro subsect. Trachygalium K. Schm. 
in Pflanzenfam. IV, 4 (1897) 151, pro sect.—Panicles lax, decussate 
at ends of stems and branches. Flowers bisexual, white. Leaves in 
whorls of 4—6, single-veined, obtuse. Mericarps glabrous, with granular 
surface. Perennial or annual herbs with very weak, retrorsely scabrous 
stems, not forming mats. 


Series 1. Palustria Pobed.—Corolla 4-fid. Stamens 4. 


34. G. palustre L. Sp. pl. (1753) 105; DC. Prodr. 4, 597; Ldb. FI. 
alt. I, 135; Ldb. Fl. Ross. II, 408; Boiss. Fl. or. III, 59; Schmalh. FI. 
II, 15; Grossh. Fl. Kavk. IV, 32; Kryl. Fl. Zap. Sib. X, 2579.—G. 
incarnatum Gilib. Fl. lith. I (1785) 2.—Ic.: Syreistsch. Fl. Mosk. gub. 
III, 192; Fedch. and Fler. Fl. 909; Rchb. Ic. Fl. Germ. XVII, tab. 
1195; Hegi, Ill. Fl. VI, 1, tab. 248.—Exs.: Fl. exs. austro-hung. No. 
2219; Fl. Sicula exs. No. 837; Fl. siles. exs. No. 828, 829 and 1082 
(under G. palustre var. maximum Moris); Fl. Cehoslov. exs. No. 85; 
Fl. stir. exs. No. 567; Pl. Finl. exs. No. 1351; Fl. Gal. and Germ. exs. 
No. 2858; Fl. Ital. exs. No. 359; Fl. Boh. and Morav. exs. No. 685; 
Pl. Finl. exs. No. 956; Fl. Pol. exs. No. 732. 

Perennial. Stems procumbent or ascending, 30-70 cm tall, 
branched, subglabrous or weakly scabrous along ribs. Leaves in distant 
whorls of 4, oblong, (8)15—20(35) mm long, (1.5)2—3(6) mm broad, 
broadened at apex, narrowed at base, single-veined, thin, terminal, 
covered with short retrorse setae along margin, or antrorse, setae along 
margin, and retrorse along middle vein; floral leaves paired, 
considerably smaller than cauline, (2)4—-5(8) mm long, 0.5—1 mm broad. 
Cymes few-flowered, terminal on stems and branches, forming lax, 
spreading, paniculate inflorescence. Peduncles doubly-triply branched, 
long, considerably longer than leaves, scabrous; pedicels equaling or 
somewhat longer than flowers, glabrous. Corolla white, 3—4 mm across, 
up to 2/3 incised into ovate, obtuse lobes. Anthers yellow. Style bifid 


332 


314 


only at tip. Fruits binate, 2 mm long, 3 mm broad, mericarps globose, 
glabrous, microgranular-tuberculate. June to August. 

In bogs, wet mossy areas and willow beds, wet marshy meadows, 
woods along rivers, subalpine meadows.— European USSR: All regions 
besides Crimea. Caucasus: All regions. Western Siberia: All regions 
(rarely in Altai). Eastern Siberia: Yenisei, Angara-Sayan. Soviet Central 
Asia: Aralo-Caspian, Balkhash Region, Syr Darya (very rarely, 
introduced). General distribution: Scandinavia, Central and Atlantic 
Europe, western Mediterranean, Balkans— Asia Minor, North America. 
Described from Europe. Type in London. 

Note. Caucasian plants of this species from subalpine meadows 
are noteworthy for their tiny leaves and flowers, and glabrous or 
subglabrous stem (var. subalpinum Pobed.). G. palustre L. is typical 
a European species, but it has spread into Asia up to Baikal. 


35. G. saturejifolium Trev. in Mag. d. Ges. naturforsch. Fl. in 
Berl. VII (1815) 146; M.B. FI. taur.-Cauc. IH, 106; DC. Prodr. IV, 
597; Ldb. Fl. Ross. II, 409, p.p.—G. palustre M.B. FI. taur.-cauc. I 
(1808) 103, non L.—G. palustre L. B. elongatum Schmalh. FI. II 
(1897) 15, p.p. 

Perennial. Stems 50—80 cm tall, straight, strong, with long branches 
all along its length, at base almost terete, above 4-angled, scabrous 
along ribs due to dense, short, laterally directed setae. Leaves in whorls 
of 4, lower ones equal, in branches somewhat unequal, oblong- 
lanceolate or linear, (20)25(30) mm long, (1.5)2(3) mm broad, acute, 
cuneate at base, narrowed into short petiole, slightly revolute along 
margin, single-veined, compact, on upper surface, along vein scabrous 
due to very short antrorse setae beneath and margin, floral leaves 
paired or solitary, considerably smaller than cauline leaves, 2-4 mm 
long, 1.5 mm broad. Inflorescence paniculate, terminal on stem and 
upper branches, ovate or globose, more or less aggregated. Peduncles 
doubly-triply- ternately branched, ovate or globose, more or less 
aggregated. Peduncles doubly-triply ternately branched, strong, highly 
scabrous; pedicels 2 times as long as flowers, glabrous or subglabrous. 
Corolla white, 2 mm across, with ovate lobes, with long cusp at tip. 
Stamens yellow. Style bipartite in upper third. Fruits binate, 1.5 mm 
long, 1.5 mm broad, greenish, glabrous, pitted; mericarps compactly 
overlapping each other along lateral walls (Plate XVIII, fig. 4). 

Along banks of rivers and on shores.— European USSR: Lower 
Volga. Caucasus: Dagestan. Endemic. Described from Astrakhan. Type 
in Bonn. 


333 


315 


36. G. karakulense Pobed. sp. n. in Addenda XXII, 712.—G. 
saturejaefolium auct. fl. As. med. non Trev.—G. elongatum auct. fl. 
As. med. non Pres}. 

Perennial. Root slender, vertical, long. Stem 40-70 cm tall, thick, 
branched, almost from the very base with spreading branches, 4-angled, 
highly retrorsely scabrous along prominent, whitish ribs. Leaves on 
entire plant in whorls of 4, oblong-lanceolate, (15)25(30) mm long, 
(3)5-7(14) mm broad, obtuse or almost rounded at apex, narrowed 
into short petiole at base or subsessile, with one prominent vein, slightly 
revolute along margin, densely retrorsely spinulose along margin, as 
well as along middle vein beneath, upper surface dark green, lower 
pale green; cauline leaves equal, leaves toward top and on branches 
unequal; floral leaves paired and solitary, tiny, (3)4(7) mm long, 
(1)2(5.5) mm broad. Cymes 5-9 flowered, in threes, terminal on stems 
and numerous branches (from very base of stem) and in ones and twos 
in axils of upper leaf whorls, tripartite, each branch with one—three 
flowers. Peduncles 3-4 cm long, scabrous, their branches horizontally 
divaricate in fruit. Pedicels short, equaling flowers or somewhat longer, 
scabrous. Corolla white, with very short tube or almost without tube, 
3.5-4 mm across, with oblong-ovate lobes, short-pubescent inside. 
Anthers brown. Style bifid at tip. Fruits mostly binate, 1.5-2 mm 
long, 3—3.5 mm broad; mericarps subglobose, finely rugose, glabrous. 
May to August (Plate XVIII, fig. 1). 

Along wet boggy banks of rivers and lakes.—Soviet Central Asia: 
Balkhash Region, Pamiro-Alai, Tien Shan. Described from Lake Kara- 
Kul (near Novo-Troitsk on Chu River). Type in Leningrad. 

Note. Leaves in all plants of this species are in whorls of four, 
with the exception of one specimen from Trans-Ili Ala Tau, which has 
the middle cauline leaves in whorls of six. 

37. G. elongatum Presl. Delic. Prag. (1822) 119; Boiss. FI. or. 
III, 59; Grossh. Fl. Kavk. IV, 32; Vizn. roslin URSR, 311.—G. palustre 
L. B. constrictum Dub. in DC. Prodr. IV (1830) 597.—G. 
saturejaefolium Ldb. Fl. Ross. II (1846-1848) 409, p.p. non Trev.— 
G. palustre B. elongatum Schmalh. FI. II (1897) 16, p.p.—Ic.: Rchb. 
Ic. Fl. Germ. XVII, tab. 1195.—Exs.: Fl. Gal. and Germ. exs. No. 
1684, 1684 bis, ter.; Fl. Palaest. exs. No. 378, Fl. Ital. exs. No. 360, 
360 bis. 

Perennial. Root slender, vertical, brown-red. Stem straight, 50- 
120 cm tall, thick, swollen at nodes, branched in upper half, with 
divaricate branches, 4-angled, with sharply prominent, whitish ribs, 
smooth, glabrous, rarely weakly scabrous. Middle leaves in whorls of 
6, equal, upper and lower leaves and those on branches in whorls of 
4, unequal, oblong-lanceolate, (12)20(25) mm long, (1.5)3(5) mm 


334 


316 


broad, obtuse, subsessile, glabrous, scabrous along margin due to sparse 
retrorse spinules, single-veined; floral leaves in whorls of 4, paired or 
solitary, (4)7(10) mm long, (1)1.5(2.5) mm broad. Inflorescence 
paniculate, terminal on stems and branches, spreading, few-flowered. 
Cymes on doubly ternately branched peduncles, the latter longer than 
leaves, scabrous. Pedicels equaling or somewhat longer than flowers, 
smooth. Corolla white with very short tube, or without tube, 3-3.5 
mm across, with ovate, acute lobes. Anthers yellow. Stamens with 
filaments arcuate within flower. Style bifid at tip. Fruits 1.5 mm long, 
2-—2.5 mm broad, with two or often with one mericarp, subglobose, 
with weakly granular-tuberculate, often rugose, glabrous. May to 
October. 

In wet meadows, along river banks.—European USSR: Upper 
Dniester. Caucasus: Western and eastern Transcaucasia, Talysh. 
General distribution: Central and Atlantic Europe, western and eastern 
Mediterranean, Balkans-Asia Minor. Described from Sicily. Type in 
Prague. 

Note. We mention reports for the Carpathians on the basis of 
literature sources (Vizn. roslin URSR). 


38. G. krymense Pobed. sp. n. in Addenda XXII, 713.—G. 
elongatum auct. fl. Taur. non Presl. 

Perennial. Root thin, vertical, with numerous, very slender branches 
from the root collar. Stem straight, 25-60 cm tall, slender, branched, 
with more or less short branches, 4-angled, weakly retrorsely scabrous, 
rooting at lower nodes. Middle cauline leaves in whorls of 6, equal, 
lower, upper and on branches in whorls of 4, unequal, narrow, linear, 
(7)20(30) mm long, (1)2(3) mm broad, obtuse, scarcely narrowed 
toward base, almost sessile, glabrous, along margin slightly revolute 
and scabrous due to very short antrorse spinules, single-veined, floral 
leaves paired, uppermost equal, tiny, (5)6(7) mm long, (0.5)1(1.5) 
mm broad. Cymes lax, 6-12 flowered, terminal on stem and branches 
forming spreading, lax, panicles. Peduncles ternately branched, long. 
Pedicels equaling flowers or 2 times as long; pedicels and peduncles 
glabrous. Corolla white, 4-5 mm across, with short tube and ovate, 
acute lobes. Anthers yellow. Stamens with filaments arcuate within 
flower. Style bifid at tip. Fruits 1.5 mm long, 2.5-3 mm broad, with 
2 globose, densely granular-tuberculate mericarps. June to August. - 

In marshes.— European USSR: Crimea. Caucasus: Ciscaucasia. 
Described from Crimea (Karagol Marsh, north of Yalta). Type in 
Leningrad. 


335 


317 


Series 2. Trifida Pobed.—Corolla trifid. Stamens 3. 


39. G. trifidum L. Sp. pl. (1753) 105; DC. Prodr. IV, 597 p.p.; 
Ldb. Fl. Ross. HI, 409, p.p.; Schmalh. Fl. II, 16, D-P.; Kryl. Fl. Zap. 
Sib. X, 2580, p.p. 

Perennial. Root slender, weak, branched. Stems decumbent or 
ascending, 15-50 cm tall, branched, glabrous or weakly retrorsely 
scabrous. Leaves in whorls of 4, linear, broad, (10)15-17(22) mm 
long, (1.5)4-5(6.5) mm broad, obtuse or rounded, gradually tapered 
toward base, thin, single-veined, glabrous on both surfaces or weakly 
scabrous along margin and beneath; floral leaves in threes or paired; 
but in latter case unequal, one smaller than other or in isolation, smaller, 
(3)6(9) mm long, (1.5)2(3) mm broad, glabrous or weakly scabrous. 
Cymes with solitary flowers or 3-flowered in axils of upper leaves. 
Pedicels and peduncles more or less long, glabrous or weakly scabrous. 
Corolla white, longer than ovary, 2 mm across, with 3 broadly ovate, 
obtuse lobes. Stamens 3. Style bifid at tip. Fruits binate, 1 mm long, 
3 mm broad. Mericarps separated, contiguous only at base, globose, 
glabrous, microgranular-tuberculate. June to August. 

In marshes and flood plains of rivers: near hot springs.— Soviet 
Far East: Kamchatka, Sakhalin. General distribution: North America. 
Described from Caucasus. Type in London. 


40. G. ruprechtii Pobed. sp. n. in Addenda XXII,-713.—G. 
trifidum var. europaeum Rupr. Fl. Samojed. (1846) 38; Fl. ingr. 496.— 
G. trifidum auct. non L.: Ldb. FI. alt. (1829) 133; Fl. Ross. II, 409; 
Weinm. Fl. Petropol. 17; Schmalh. FI. II, p.p. 6.—Ic.: Syreistsch. FI. 
Mosk. gub. III, 193.—Exs.: Pl. Finl. exs. No. 361 a and b, No. 1352; 
GRF, No. 1477. 

Annual. Root extremely slender, filiform, creeping. Stems 
numerous, very slender, lodged, 10-30 cm tall, many-branched, 4- 
angled, retrorsely scabrous along ribs, with crowded, short, retrorse 
setae. Leaves in whorls of 4, of which 2 somewhat, or considerably, 
shorter than other two, oblong-oblanceolate, (5)10(15) mm long, 
(1)1.5(2.5) mm broad, obtuse and somewhat broadened at apex, 
gradually narrowed at base, single-veined, thin, on both surfaces green, 
retrorsely scabrous along margin and along vein beneath, rarely 
glabrous; floral leaves of same shape and pubescence, but considerably 
smaller than cauline, (1)3(6) mm long, 0.5-1 mm broad, in threes, 
paired (here one somewhat smaller than other) or isolated. Cymes 
terminal on stems and branches and axillary, 1-2, rarely 3-flowered, 
peduncles considerably longer than leaves, together with pedicels, 
retrorsely scabrous. Pedicels 5-10 times as long as flowers, 


336 


318 


(4)8-10(15) mm long, horizontally inclined or arched downward and 
considerably elongated in fruit. Corolla white, 1 mm across, equaling 
or shorter than ovary, with suborbicular lobes. Stamens 3. Anthers 
yellow. Style bifid at tip. Fruits 1 mm long, 2 mm broad. Mericarps 
globose, contiguous only at base, glabrous, microgranulate. Second 
half of June to July (Plate XVIII, fig. 3). 

On peat and sedge marshes, banks of springs.— European USSR: 
Karelia-Lapland, Dvina-Pechora, Baltic Region, Ladoga-IImen, Upper 
Volga, Volga-Kama, Middle Dnieper, Volga-Don, Black Sea Region. 
Western Siberia: Ob’ Upper Tobol, Irtysh. Eastern Siberia: Yenisei, 
Dauria (rarely), Lena-Kolyma. Soviet Central Asia: Aralo-Caspian, 
Dzhungaria-Tarbagatai. General distribution: Scandinavia. Described 
from Peterburg (Port Grebnoi, Galernaya Harbor?). Type in Leningrad. 

Note. Linnaeus described the species G. trifidum from Canada; 
European and Asian plants, which are not identical to it, have been 
incorrectly referred to this species. Ruprecht was the first to notice the 
differences and named the European plants G. trifidum var. europaeum. 
Subsequently, D.I. Litvinov, while distributing exsiccatae from Pskov 
(GRF, No. 1477), indicated that the European species differs from the 
American one in scabrousness, in having very thin, low stems, very 
small corollas, equaling or shorter than the ovary, whereas in the 
American plant the corolla is longer than the ovary. This species is 
widespread, occurring all over Siberia, but it does not occur in the 
Crimea, Caucasusor, mountainous parts of Soviet Central Asia. From 
the Yenisei to the Soviet Far East, it is replaced by the closely related 
species G. baicalense. 


41. G. baicalense Pobed. sp. n. in Addenda XXII, 714.—G. 
trifidum auct. fl. Sib. non L.—G. trifidum var. brevipedunculata Rgl. 
Tent. fl. ussur. (1861) 77. 

Annual. Roots very thin. Stems one to ten, 5—20 cm tall, filiform, 
straight or lodged, ascending, branched, 4-angled, scabrous along ribs 
due to retrorse setae. Leaves in whorls of 4, elliptic, (5)7—8(11) mm 
long, (1)2(2.5) mm broad, obtuse, not broadened at apex, abruptly 
narrowed at base into extremely short petiole, glabrous or retrorsely 
scabrous along margin and middle vein beneath; floral leaves paired 
or solitary, considerably smaller, (1.5)2.5(3) mm long, 0.5—1 mm broad, 
but similar in shape and pubescence. Cymes 3-4 flowered, rarely 1- 
2 flowered in axils of upper leaves and at ends of lateral branches. 
Pedicels (1.5)3(8) mm long, equaling or somewhat longer than adjacent 
leaves, glabrous or barely retrorsely scabrous. Peduncles retrorsely 
scabrous. Corolla longer than ovary, 2 mm across, with 3-4 broadly 
ovate, obtuse lobes. Stamens 3-4. Style bifid at tip. Fruits binate, with 


319 


2 separated, globose mericarps, contiguous only at base, occasionally 
with one mericarp, other rudimentary, 1 mm long, 2 mm broad, black, 
glabrous, microgranular-tuberculate. June to August (Plate XVIII, 
fig. 2). 

On sphagnum and sedge-cotton grass marshes, mossy banks of 
rivers and streams, wet and marshy meadows.— Eastern Siberia: Lena- 
Kolyma, Angara-Sayan, Dauria. Soviet Far East: Zeya-Bureya, 
Kamchatka, Ussuri. Endemic. Described from Trans-Baikal. Type in 
Leningrad. 

Note. Differs from G. ruprechtii Pobed. by short, glabrous pedicels 
(and not long, retrorsely scabrous), elliptical (and not oblanceolate) 
leaves, corollas 2 mm across (and not 1 mm). 


Section 8. LEPTOGALIUM Lange in Willk. et Lange Prodr. fl. Hisp. 
(1870) 316.—Leiogalia DC. Prodr. IV (1830) 593, p.p., pro subsect.— 
Cymes erect, lax or gathered in umbellate panicles. Flowers bisexual. 
Leaves in whorls of 6-8, single-veined and with cusp. Fruits straight, 
glabrous, surface granular. Perennial herbs with low, glabrous, rarely 
retrorsely scabrous, straight or decumbent stem, forming mat. 


Series 1. Hercynica Pobed.—Stems smooth. 


42. G. hercynicum Weig. Observ. bot. (1772) 25; DC. Prodr. IV, 
598; Clapham, Tutin, Warburg, Fl. of Brit. isl. 933.— G. saxatile L. 
Sp. pl. (1762) 154, p.p. (solum planta Saxon.) non L. 1753, 106.— 
Ic.: Fl. Germ. XVII, tab. 1194 (subg. saxatile); Hegi, Ill. Fl. VI, 1, f. 
120.— Exs.: Fl. Boh. and Morav. exs. No. 564; Fil. siles, exs. No. 
1344; Fl. Gall. and Germ. exs. No. 2077; Pl. Finl. exs. Nos. 363, 
1353; Fl. Germ. exs. No. 2420. 

Perennial. Stems 10-30 cm tall, decumbent or ascending 4-angled, 
glabrous, branched. Leaves in whorls of 5—6, upper leaves in whorls 
of 3-4, oblanceolate, 5(6)10 mm long, 1(1.5)3 mm broad, abruptly 
chondroid-acuminate at apex, gradually attenuated at base, with short 
petioles, slightly revolute along margin, single-veined, glabrous or 
scabrous along margin due to short, antrorse setae. Cymes terminal, 
lax, 2—3-flowered or gathered in umbellate panicles. Peduncles 8-15 
mm long. Pedicels almost equaling mericarps or somewhat longer; 
pedicels and peduncles glabrous. Corolla white, 3.5—4 mm across, 
lobes ovate. Fruits binate, 1-1.5—2 mm in diameter, glabrous, granular- 
tuberculate. June to August. 

In moss, wet coniferous forests, forest edges.— European USSR: 
Ladoga-IImen (only in Karelian Isthmus). General distribution: 
Scandinavia, Central and Atlantic Europe. Described from Switzerland. 
Type in London. 


33 


(o/<) 


320 


Note. Within the boundaries of the USSR (on Karelian Isthmus) 
lies the eastern boundary of the distribution of this species. A report 
of the occurrence of the species in the Urals [Pall. Iter. I, (1771) 379] 
is evidently erroneous. 


43. G. tianschanicum M. Pop. in Byull. Mosk. obshch. ispyt. prir. 
XLVII (1938) 88.—G. tianschanicum 1. xerophytica M. Pop. and 2. 
prostrata M. Pop. |.c.—G. saxatile var. leiophyllum Trautv. in Bull. 
Soc. Nat. Mosc. XXXIX, 1 (1866) 333; O. and B. Fedch. Perech. rast. 
Turkest. III, 140.—G. saxatile auct. fl. As. med. non L.—G. 
petrophilum Pavl. in sched. 

Perennial. Rootstock branched, with slender, procumbent branches. 
Stems 5-10 cm tall, straight, decumbent or ascending, 4-angled, simple 
or branched from base, botryoid, glabrous. Leaves in whorls of 4 or 
5-6, oblong or lanceolate, 4-17 mm long, 1-5 mm broad, acute or 
subobtuse, at top abruptly chonroid acuminate, but margin slightly 
revolute, at base gradually attenuated into short petiole, fairly stiff, 
almost coriaceous, single-veined, glabrous. Cymes terminal on stems 
and branches, simple with 3 flowers. Pedicels 2 times as long as 
flowers, divaricate with 2-4 floral leaves at base, resembling cauline 
leaves but smaller. Corolla yellow (whitish yellow), 3.5 mm across 
with ovate lobes. Fruits binate, 1-1.5 mm in diameter, globose, 
microgranulate. June to August (Plate XVIII, fig. 3). 

On stony mountain slopes in subalpine zone and in detrital 
deposits.— Soviet Central Asia: Dzhungaria-Tarbagatai, Tien Shan. 
Endemic. Described from Trans-Ili Ala Tau. Type in Alma-Ata. 

Note. In describing this species the following forms were 
distinguished: 1) xerophytica M. Pop.—light green, with low, straight 
stems and small, lanceolate, acute leaves and 2) prostrata M. Pop., 
with elongated, decumbent stems, larger, oblong, subobtuse, green 
leaves. Most of Schrenk’s specimens from the Dzhungarian Ala Tau 
belong to the latter form. The first form is represented with us, 
apparently, by Semenov’s specimen from the Trans-Ili Ala Tau. 

Plants from Talas Ala Tau differ somewhat from the indicated 
forms in having greater branching and closeness of leaf whorls and 
somewhat broader leaves. N.V. Pavlov, on the herbarium labels, 
annotated them as a distinct species.—G. petrophilum Pavl., but did 
not describe it. Evidently, this form has the same rank as the two 
others named. 


44. G. anisophyllum Vill. Prosp. (1779) 20 and Hist. Pl. Dauph. 
(1787) 317 bis.; Boiss. Fl. or. III, 54; Hegi. Ill. Fl. VI, 1, 218; Vizn. 
roslin URSR, 313; Szaf. Kulcz. and Pavl. Rosliny Polsk. 619.—Ic.: 


339 


321 


Vill. Hist. Pl. Dauph. II, tab. 7; Hegi. Ill. Fl. tab. 119 e bis h.—Exs:.: 
Fl. Gal. and Germ. exs. No. 3403; Fl. stir. exs. No. 569. 

Perennial. Rootstock slender, branched. Stems one-several, 
branched from base, 5—10 cm tall, rarely 15 cm tall, 4-angled, glabrous. 
Leaves in whorls of 5—8, lower leaves and in branches sometimes in 
whorls of 4, oblanceolate-linear or oblong-obovate, 7-13 mm long, 1-— 
2.5 mm broad, with more or less long cusp at apex, gradually attenuated 
at base into short petiole, single-veined, glabrous; floral leaves paired 
or in threes, considerably smaller than cauline leaves. Cymes in threes, 
terminal on stems and branches, furcately or ternately branched, more 
rarely simple, 3—4—9-flowered, with more or less long peduncles, 
pedicels shorter than flowers, glabrous, greatly thickened at top. Flowers 
yellow-white, 3.5—4 mm across, with ovate, subobtuse lobes. Anthers 
large, broad, white-yellow. Style bipartite in upper third. Fruits with 
one mericarp, rarely binate, glabrous, 1-1.5 mm long, 2-3 mm broad, 
granular-tuberculate. June to September. 

In rocks, stones, stony screes, in alpine and subalpine zones.— 
European USSR: Upper Dniester. General distribution: Central and 
Atlantic Europe, Balkans-Asia Minor. Described from France. Type in 
France. 

Note. This apparently is a species aggregate, whose separate races 
are described as the species G. alpestre Cand., G. bocconi Haenke, G. 
sudetica Tausch. and others. The eastern Carpathians are the eastern 
boundary of its distribution. 


45. G. pumilum Murr. Prodr. Gotting. (1770) 44; Szaf. Kulcz. 
Pavl. Rosliny Polsk. (1953) 620.—G. silvestre Poll. Hist. Pl. Palat. I 
(1776) 151; Fedch. and Fler. Fl. 913.—Exs.: Fl. Ital. exs. No. 1357 
[sub G. silvestre var. austriacum Jacq. (pro sp.)]; Fl. Boh. and Morav. 
exs. No. 568 (sub G. austriacum Jacq.), No. 490 (sub G. asperum 
Schreb.); Billot Fl. Gall. and Germ. exs. No. 378 bis (sub G. silvestre 
Poll.). 

Perennial. Rootstock branched. Stems several, weak, ascending or 
lodged, with fruit-bearing and sterile shoots, 10-50 cm tall, 4-angled, 
smooth, not blackened when dry. Leaves in whorls of 6-8, linear- 
oblanceolate or linear, cuspidate, slightly revolute and scabrous along 
margin due to retrorse setae; elsewhere more or less glabrous, single- 
veined, basal leaves smaller. Cymes terminal on stems and branches, 
forming almost corymbose panicle on long peduncles, longer than stem 
internodes. Pedicels short, twice as long as flowers. Corolla cream 
colored, 2.5—4 mm across with acute, but not mucronate lobes. Fruits 
binate, 1 mm long, covered with obtuse tubercles. June to August. 

In dry light forests, in clearings.— European USSR: Baltic Region 
(only in Kaliningrad Region). General distribution: Central and Atlantic 


340 


322 


Europe, western Mediterranean, Balkans-Asia Minor. Described from 
Gottingen. Type in Berlin. 

Note. This species is included on the basis of the literature 
[Abromeit, Fl. Ost. u. Westpreussen (1940) 362], since there is no 
herbarium material within the territory of the USSR except for the 
single specimen collected by Kropachev on the southern bank of Lake 
Ladoga, where the occurrence of this species is extremely doubtful. 
Extension of its range into the Kaliningrad Region is entirely probable, 
since it is distributed in the adjoining regions of Poland. 


46. G. vartanii Grossh. in Tr. po geobot. obsled. pastbishch. Azerb. 
vyp. 1 (1929) 78 and Fl. Kavk. IV, 33.—Ic.: Grossh. I.c. (1929), tabl. 
VIII. 

Perennial. Rootstock vertical, slender, woody. Stems several from 
root, 5-15 mm tall, weakly branched, 4-angled, glabrous, sterile shoots 
densely foliated, more slender. Stems bearing inflorescence, with 
spreading leaf whorls. Leaves in whorls of 6, flat, thicker, narrowly 
linear or oblanceolate-linear, (4)10(15) mm long, 1—1.5 mm broad, 
narrowed at both ends, with cusp at apex, with short, densely pubescent 
petiole, single-veined, glabrous on both surfaces. Floral leaves 6-2-1 
in number, considerably smaller than cauline leaves, 3-4 mm long, 
0.5-1.5 mm broad. Cymes terminal, 6—7-flowered. Peduncles ternately 
or furcately branched, together with pedicels thicker, glabrous. Pedicels 
equaling flowers or somewhat longer, thickened at top. Corolla 6-7 
mm across, with oblong obtuse lobes, united at base into a very short 
tube. Anthers yellow, large. Style bipartite almost from base. Fruits 
immature, young—minute, 1 mm long, 1.5 mm broad, dark brown, 
surface microgranulate. June to July. 

On rubbly mountain slopes in alpine zone.— Caucasus: Eastern 
Transcaucasia. Endemic. Described from Nukha. Type in Baku. 

Note. This species was erroneously placed in section Eugalium, 
whereas its actual position is in section Leptogalium Lge. in a series 
of vicariant species: G. hercynicum Weig. on the Karelian Isthmus, G. 
tianschanicum M. Pop. in Central Asia, G. vartanii Grossh. in the 
Greater Caucasus Range, G. kiapazi Mand., in the eastern Caucasus. 
In the mountains of Western Europe there is a series of vicariant races 
with diverse, disjunct ranges. 


47. G. kiapazi Manden. in Zam po sist. i. geogr. rast. vyp. 17 
(1953) 64. 

Perennial. Rootstock slender, filiform, creeping, woody, brown. 
Stems several from root, much-branched with divaricate branches, 5— 
8 cm tall, 4-angled, glabrous, sterile and flowering shoots densely 


341 


323 


foliated. Leaves in whorls of 6, flat, thicker, oblanceolate or oblong- 
lanceolate, upper leaves and those in branches, often almost elliptic, 
4—6 mm long, 1.5—2 mm broad, with cusp at apex, somewhat narrowed 
and weakly pubescent toward base, single-veined, glabrous on both 
surfaces; upper leaves in whorls of 6 or paired, slightly smaller than 
cauline leaves. Cymes simple; axillary cymes 1-flowered, and terminal 
3-flowered, on long pedicels, 2-3 times as long as than flower (but 
not longer than floral leaves), thicker, especially thickened at tip, 
glabrous. Corolla white, 3-4 mm across, with ovate, short-acuminate 
lobes. Anthers yellow. Style bipartite up to 1/3. Fruits binate, immature, 
young 1 mm long, 0.75 mm broad, dark brown, microgranulate. July 
to August. 

On stony slopes of alpine zone.— Caucasus: Eastern Transcaucasia 
(southern part). Endemic. Described from Mt. Kyapaz. Type in 
Leningrad. 

Note. This species is very close to G. vartanii Grossh. and differs 
in general appearance (three-flowered terminal cymes and single- 
flowered axillary ones, with pedicels no longer than the floral bracts); 
the leaves of G. kiapazi are somewhat broader than the leaves of G. 
vartanil. 


Series 2. Uliginosa Pobed.— Stems retrorsely scabrous along ribs. 


48. G. uliginosum L. Sp. pl. (1753) 106; DC. Prodr. IV, 597; 
Schult. Syst. veg. III, 247; Besser, Enum. pl. Volh. 7; Bge. in Ldb. FI. 
alt. 1, 135; Fl. Ross. III, 408; Grossh. Fl. Kavk. IV, 33; Kryl. Fl. Zap. 
Sib. X, 2578.—G. grandiflorum Gilib. Fl. lith. I (1785) 4.—Ic.: 
Syreistsch. Fl. Mosk. gub. II, 191; Rchb. Ic. Fl. Germ. XVII, tab. 
1193, 1194; Hegi, Ill. Fl. VI, 1, tab. 248, 4.—Exs.: Pl. Finl. exs. No. 
955; Fl. Pol. exs. No. 731; Fl. Boh. and Morav. exs. No. 565; FI. siles. 
exs. No. 37, No. 827; Fl. Gall. and Germ. exs. No. 2076. 

Perennial. Stems spreading, weak, 15-70 cm tall, branched, 
clinging because of short, thin, retrorse spinules along ribs. Leaves in 
whorls of 6, oblanceolate or lanceolate, (3)10(16) mm long, (1)1.5(3) 
mm broad, with more or less long cusp at apex, gradually narrowed 
toward base, single-veined, barely revolute along margin, with thin, in 
portions retrorse spinules along margin as well as vein beneath; floral 
leaves paired, uppermost leaves solitary, very small, (1)2(4) mm long, 
(0.25)0.5(1) mm broad. Cymes lax, 6-9-flowered terminal on stems 
and branches, forming paniculate, spreading inflorescence. Peduncles 
ternately branched, long. Pedicels shorter than or as long as flowers; 
pedicels and peduncles clinging due to spinules. Corolla white, 3—3.5 
mm across, united up to % the length from base into short, broad tube, 


324 


with ovate, acute, lobes. Anthers yellow. Style bipartite in upper third, 
with 2 fleshy bilobed nectaries at base. Fruits 1 mm long, 1.5—2 mm 
broad, binate, rarely with one mericarp, another rudimentary, densely 
finely tuberculate. June to September. 

Along margins of peat bogs, canals, on wet banks, wet meadows.— 
European USSR: Karelia-Lapland, Dvina-Pechora, Baltic Region, 
Ladoga-IImen, Upper Volga, Volga-Kama, Upper Dnieper, Middle 
Dnieper, Volga-Don, Black Sea Region. Caucasus: Ciscaucasia, western 
(southern part) and eastern Transcaucasia. Western Siberia: All regions. 
Eastern Siberia: All regions. Soviet Central Asia: Aralo-Caspian. 
General distribution: Arctic (Greenland) Region, Scandinavia, Central 
and Atlantic Europe, Mongolia. Described from Western Europe. Type 
in London. 

Note. Western Siberian plants differ by having taller stature and 
larger leaves (var. sibiricum m.). In the Far East, this species is replaced 
by G. dahuricum Turcz. 

This species sometimes occurs also in upland meadows in the 
form of extremely small plants with very small leaves and flowers (an 
ecological form). 


49. G. dahuricum Turcz. Fl. baic.-dah. I (1842-1845) 530; Ldb. 
Fl. Ross. II, 1, 409; Maxim. Prim. Fl. amur. 140; Rgl. Tent. Fl. ussur. 
77; Kom. in Tr. SPb. bot. sada, XXV, 492; Kom. and Alis. Opred. 
rast. Dalnevost. kr. II, 961.—G. asperulum a. typicum and B. 
dahuricum Maxim. in Mél. Biol. IX (1877) 262.—G. triflorum Korsh. 
in Tr. SPb. bot. sada, XII (1893) 347, No. 275, non Michx .—Ic:: 
Kom. and Allis. l.c., tabl. 287; Herder in Bull. Soc. Nat. Mosc. XX XVII, 
tab. 3. : 
Perennial. Rootstock slender, creeping, with numerous shoots. 
Stems weak, 30-50 cm tall, ascending, branched, sharply 4-angled, 
retrorsely scabrous along ribs due to scattered spinules. Leaves in 
whorls of 5—6, obovate-oblong, (11)20(40) mm long, (2)5—7(9) mm 
broad, abruptly short-acuminate at apex, gradually attenuated toward 
base into short petioles, single-veined, with scattered retrorse setae 
along margin and sometimes along veins beneath; floral leaves in whorls 
of 2-4 or solitary, (3)5(7) mm long, (1)1.5(2.5) mm broad. Cymes 
terminal and in axils of upper leaves, simple, spreading, 2—3-flowered. 
Peduncles and pedicels trichoid, long, 2-4 times as long as flowers, 
glabrous. Corolla 3-4 mm across, with broadly ovate, short-acuminate 
lobes. Anthers yellow. Style bipartite in upper 1/3 part. Fruit 2 mm 
long, 1.5—2 mm broad, binate or with one mericarp, second rudimentary, 
342 oblong, almost black, glabrous, tuberculate or pubescent with more or 


345 


325 


less long, shiny hairs. Fruits in this species only a few, sometimes a 
solitary fruit may be seen in the entire plant. June to August. 

In meadows, scrub thickets; marshy meadows, willows, forest 
clearings.— Eastern Siberia: Lena-Kolyma, Angara-Sayan, Dauria. 
Soviet Far East: Zeya-Bureya, Uda Ussuri. General distribution: Japan, 
China. Described from Trans-Baikal, from Argun River. Type in 
Leningrad. 

Note. Forms with pubescent and glabrous fruits are geographically 
differentiated: in Eastern Siberia only the glabrous form is seen; 
whereas in the Soviet Far East, it is found along with the pubescent 
form. 

In Yakutia, a form with large flowers (var. grandiflorum m.) grows. 
In North America it is replaced by the widespread species G. asprellum 
Michx. (described from Canada), which Maximovicz believed to be 
identical with G. dahuricum, but, as quite correctly indicated by V.L. 
Komarov (Fl. Manch. 492), both species are well-distinguished by the 
following characters: G. dahuricum Turcz. has obovate (and not 
lanceolate) leaves and petals twice as large; the fruits may be glabrous 
as in G. asprellum or pubescent. The habit of the eastern Asian species 
with considerably less-branched stems and larger leaves sharply 
distinguishes it from the American species. 


50. G. vassilezenkoi Pobed. sp. n. in Addenda XXII, 741. 

Perennial. Stem thick, up to 100 cm long, with long divaricate 
branches, succulent, flattened when dry, weak, lodged, 4-angled, 
lustrous, retrorsely scabrous along ribs. Leaves in whorls of 5—6; upper 
leaves in whorls of 4, elliptic or oblong-elliptic, (20)30(40) mm long, 
(7)10(14) broad, rounded at apex with short thin mucro, gradually 
narrowed at base into short petiole or nearly sessile, with one prominent 
vein, thin, with short antrorse or often retrorse spinules along margin 
and vein beneath; floral leaves paired and solitary, very small, (2)2.5- 
3(5) mm long, (0.5)1(2.5) mm broad. Cymes terminal on stems and 
lateral branches, 7—9-flowered, on long, scabrous, furcately or ternately 
branched peduncles. Pedicels 2—3 times as long as flowers, rarely 
almost as long, glabrous or scabrous. Corolla considerably larger than 
ovary, with short, broad tube, 3.5 mm across, with ovate, acute lobes. 
Anthers brownish, large, filaments curved within flower. Style bipartite 
almost from base and with 2 fleshy semilunar large lobes of nectaries. 
Fruits binate, 2 mm long, 4 mm broad, mericarps globose, densely 
covered with hooked hairs. June to August. 

In rocky ravines, on banks of rivers and lakes.— Soviet Central 
Asia: Pamiro-Alai. Described from Sangardak River. Type in Leningrad. 


346 


326 


51. G. pojarkovae Pobed. sp. n. in Addenda XXII, 715. 

Perennial. Root more or less woody, gray-brown. Stems several 
from root, weak, lodged, 10—60 cm tall, 4-angled, retrorsely scabrous 
along ribs, branched. Leaves in whorls of 6, thin, oblanceolate or 
elliptic, (10)20—25(30) mm long, (2)5(6) mm broad, gradually narrowed 
at apex and abruptly short-acuminate, attenuated at base into hardly 
noticeable petiole or almost sessile, single-veined, with tiny, antrorse 
spinules along margin, otherwise glabrous; floral leaves in whorls of 
3-5, uppermost paired or solitary, gradually becoming smaller from 
bottom upward, (3)6(7) mm long, 1-3 mm broad. Cymes few-flowered, 
aggregated at ends of stems and branches and in axils of upper whorls 
in lax, small panicles. Peduncles doubly, ternately or furcately 
branched. Corolla 5-6 mm across, with ovate, obtuse lobes. Anthers 
large, yellow, filaments inserted just below margin of corolla tube. 
Style bipartite from base, emerging from center of surrounding discoid 
nectary. Fruits binate, with equal mericarps or with one smaller than 
other or with reduced second mericarp 2 mm long, 4 mm broad. 
Mericarps globose, with granular surface, sparsely covered with short, 
hooked, lamellar, white, shiny, hairs. June to July. 

In alpine zone along banks of springs and meltwater pools.— 
Soviet Central Asia: mountainous Turkmenia. Endemic. Described from 
central Kopet Dag. Type in Leningrad. 


Section 9. PLATYGALIA DC. Prodr. (1830) 598. pro subsect. p.p.; 
Lange in Willk. and Lange Prodr. fl. Hisp. II, 308.—Panicles terminal, 
many-flowered. Flowers bisexual. Leaves in whorls of 4, with three 
distinctly prominent veins, without mucro at apex. Fruits glabrous or 
hamosely setulose. Perennial herbs with straight, more or less sturdy 
stems, not forming mats. 


Series 1. Rubioidea Pobed.—Fruits rugose, with pericarp loosely 
surrounding the seeds, usually entirely glabrous. Leaves large, broad. 


52. G. articulatum Lam. Ill. I (1791) 260; DC. Prodr. IV, 509.— 
[non Valantia articulata L. Sp. pl. (1753) 1052]—G. geniculatum 
Roem. and Schult. Syst. veg. III (1818) 215; Grossh. Fl. Kavk. IV, 
31; Vizn. roslin URSR, 312.—G. rubioides M.B. FI. taur.-cauc. I (1808) 
102; DC. 1.c. 599, p.p.; Boiss. Fl. or. III, 48; Vizn. roslin URSR, 312, 
p.p.—G. rubioides B. leiocarpum Choroschk. in Syreistsch. Fl. Mosk. 
gub. I (1906) 198.—G. rubioides var. eriophyllum Bordz. in Zap. 
Kievsk. bot. sada, V—-VI (1927) 40 and XII-XIII (1931) 36; Fedde, 
Repert. 30, 393.—G. boreale B. rubioides Schmalh. Fl. II (1897) 16, 
p-p.—G. boreale b. geniculatum Roem. and Schult. (sp.) l.c.— 


343 


327 


Plate XIX. 
1. Galium volgense Pobed., upper part of plant, corolla, fruit with one mericarp 
in cross section and other entire.—2. G. articulatum Lam., upper part of plant 
and fruit with one mericarp in section —3. G. amblyophyllum Schrenk, upper 
part of plant, corolla, fruit—4. G. valantioides M.B., upper part of plant, 
corolla, fruit. 


347 


328 


‘G. physocarpum Chrsh. in Vizn. roslin URSR (1950) 312, non Ldb.— 


Exs.: Pl. Abch. exs. No. 623. 

Perennial. Rootstock creeping, reddish, with one—three stems. Stems 
30-100 cm tall, straight or ascending at base, stout, scabrous along 
ribs due to dense, hard antrorse setae, often with soft, short down 
along edges, especially on lower internodes. Leaves in whorls of 4, 
broad, oblong-ovate or elliptic, (2)5.5—6.5(11.5) cm long, (0.6)1—2(3.5) 
cm broad, obtuse or subobtuse, with three, sometimes 5 sharply 
prominent veins, upper surface glabrous, lower and middle leaves with 
soft short hairs over entire surface beneath or, like upper ones, with 
antrorse setaceous hairs along veins and margin, thick, rigid, with 
short petioles; floral leaves paired, toward inflorescence solitary, small, 
(3)5—6(10) mm long, (1)1.5—2(3) mm broad, broadly ovate or elliptic, 
obtuse, sessile, together with peduncles and pedicels shortly, setosely 
pubescent over entire surface, or only beneath, or only along margin, 
rarely entirely glabrous. Panicle dense, broad, dichotomously branched, 
at oneset of flowering short, compact, subsequently almost corymbose, 
cuneate. Cymes 3-4, each 30-40-flowered, aggregated on ends of 
stems, apical peduncles short, doubly furcately and ternately branched, 
often with 4 branches, lateral peduncles longer. Pedicels equaling 
flowers or barely longer. Corolla 3—3.5 mm across, yellowish, with 
broadly elliptic obtuse lobes. Fruits glabrous, 1.5 mm long, 2.5-3 mm 
broad, coarsely rugose because of drooping walls of pericarp. Fruit set 
is never complete in this species, only a few occur in each individual 
plant. May to September (Plate XIX, fig. 2). 

In wet meadows, scrubs, along banks of rivers and seas, on flood- 
plains, in forest edges on mountain slopes.— European USSR: Ladoga- 
Ilmen (introduced), Upper Dnieper, Middle Dnieper, Volga-Don, 
Bessarabia, Black Sea Region, Crimea, Lower Don, Lower Volga. 
Caucasus: Ciscaucasia, Dagestan, western, eastern and southern 
Transcaucasia. Endemic. Described from the ‘Near East’. Type in 
London. 

Note. Roemer and Schultes reported the distribution of this species 
as “In Oriente” (In the East), thereby implying the southern part of the 
Caucasus. Lamarck’s report from the East (Levant) referred to cultivated 
plants. 

This species easily hybridizes not only with species of this series 
(G. volgense Pobed.) but also with G. boreale L., which belongs to a 
separate group of species, comprising many hybrid forms, which 
approach one or the other parent species. Sometimes hybrids of this 
species were described as separate forms; for example, Khoroshkov 
described G. rubioides B. trichocarpum Choroschk. This species is 
well-distinguished from the European G. rubioides L. not only in the 


329 


characters described by its authors, but also in the presence of soft, 
short pubescence on the lower. or sometimes also on the upper surface 
of the lower and middle leaves. Such pubescence is never found on 
the Central European plants. 

G. physocarpum Ldb., described from Astrakhan [Fl. Ross. II 
(1844) 410], differs only in having greatly swollen fruits, 2.5-3 mm 
long and 3-4 mm broad. When it was established that a plant with 
such fruits was occasionally seen along the Don, around Kiev and in 
other places, and that the swollen fruit is a pathological phenomenon 
caused by insects, G. physocarpum stopped being treated as a species 
and was given the rank of variety or form. However, in the Volga delta 
and in Krasnoarmeisk (formerly Sarepta) almost all plants resembling 
G. articulatum, possess large, swollen fruits, and, hence, a special 
anatomical investigation is required. 

Economic importance. On account of its large, dense panicles of 
white flowers and broad, stiff leaves, this species is often grown as a 
decorative plant in gardens and parks and easily escapes, even in an 
environment not natural to its original location: in the Leningrad 
Botanical Garden, in European gardens in the northern part of Central 
Europe, etc. The plant is easily brought in with the seeds of other 
cultivated plants. 


53. G. rubioides L. Sp. pl. (1753) 105; Scop. Fl. Carn. I, 100; 
DC. Prodr. IV, 599, p.p.; Vizn. roslin URSR, 312, p.p.; Schmalh. FI. 
II, 16, p.p.—Ic.: Schl. Lang and Schenk, Fl. Deutschl. Ed. 5, XXVIII, 
2892; Rchb. Ic. Fl. Germ. XVII, tab. 1186; Hegi, Ill. Fl. VI, 1, 223.— 
Exs.: Fl. exs. austro-hung. No. 2201. 

Perennial. Rootstock creeping, reddish, with one-three stems. Stems 
40-80 cm tall, straight or ascending, stout, scabrous along ribs in 
lower and middle part due to dense, hard, antrorse setae, glabrous in 
upper part and along edges, with a ring of short hairs at nodes at base 
of leaves, branched almost over its entire length, with long, antrorse 
branches. Leaves in whorls of 4, ovate-oblong, ovate or elliptic, (2.5)4— 
4.5(5) cm long, 0.8—1.5 cm broad, 3-veined, subobtuse, short petioles, 
scabrous due to short, hard, antrorse setae along margin and veins 
beneath, less rigid than in preceding species; floral leaves paired, 
upper leaves single, (3)4(5) mm long, (1.5)2(2.5) mm broad, broadly 
elliptic, rounded at apex, fimbriate or entirely glabrous. Panicle at 
onset of flowering more lax than in preceding species, subsequently 
pyramidal, dichotomously branched. Peduncles doubly furcately and 
ternately branched, 25—30-flowered, together with pedicels glabrous 

348 or with few, scattered, setaceous hairs. Corolla white, 3.5—-5 mm across, 


330 


with elliptic lobes, abruptly narrowed at tip. Fruits glabrous, 1.5—2 
mm long, 2—2.5-3 mm broad, rugose. June to August. 

Along river banks, in meadows, in forest glades and edges.— 
European USSR: Upper Dniester. General distribution: Central Europe 
(southern part). Described from Central Europe. Type in London. 


54. G. volgense Pobed. sp. n. in Addenda XXII, 715.—G. boreale 
B. rubioides Schmalh. Fl. II (1897) 16, p.p.—G. rubioides auct. fl. 
Ross. non L.—G. rubioides var. subphysocarpum Korsh. in Bull. Acad. 
Sc. Pétersb. V, ser. 1 (1894) 103.—G. hyssopifolium Chrshan. in Vizn. 
roslin URSR (1949) 312, non Hoffm. 

Perennial. Rootstock slender, creeping. Stems 25-80 cm tall, 4-6 
in number, ascending or straight, slender, simple or branched, at base 
glabrous or scabrous along ribs due to short, hard, antrorse setae. 
Leaves lanceolate, (1.5)5.5-6.5(7.5) cm long, (0.5)0.7-0.8(1.8) cm 
broad, gradually attenuated at apex, acute, rounded at base, 3-veined, 
thin, soft, lower surface lighter, glabrous on both surfaces or scabrous 
along veins beneath and margin due to short, antrorse setae, sessile or 
subsessile, connate at base of whorl into short pubescent ring; floral 
leaves small, 5-7 mm long, 1.5-4 mm broad, ovate, broadly ovate, 
broadly elliptic, obtuse, sessile, glabrous. Panicle more or less lax, 
cuneate and in axils of upper leaves, dichotomously branched, more or 
less long, spreading or slightly aggregated. Cymes 7—15-flowered. 
Peduncles ternately or doubly ternately branched, glabrous together 
with pedicels. Pedicels equaling flowers or somewhat longer. Corolla 
white, 4-5 mm across, with oblong, acute or subobtuse lobes. Fruits 
glabrous, 1.5-2 mm long, 2-3 mm broad, rugose; pericarps loosely 
surrounding seeds, initially greenish, on maturing brownish-dark purple, 
very. often swollen, 2 mm long, 2.5 mm broad. June to September 
(Plate XIX, fig. 1). 

In meadows, scrub, wet shady forests, along riverbanks.—European 
USSR: Dvina-Pechora, Ladoga-IImen, Upper Volga, Volga-Kama, 
Upper Dnieper, Middle Dnieper, Volga-Don, Volga Region, Bessarabia, 
Black Sea Region, Lower Don. Western Siberia: All regions. Soviet 
Central Asia: Aralo-Caspian, Dzhungaria-Tarbagatai. Endemic. 
Described from Volga River (village of Uzmorye). Type in Leningrad. 

Note. Hybrids of this species with G. boreale occur in the northern 
European part of the USSR and in Western Siberia; besides, in habit 
the plant resembles G. volgense, but the fruits are sparsely pubescent. 


55. G. ussuriense Pobed. in Addenda XXII, 716.—G. rubioides 
auct. fl. Orienti Extremi. 


331 


349 Perennial. Stem 50-90 cm tall, stout, straight, with shortened, sterile 
branches, glabrous along ribs and edges, with ring of dense, short 
hairs only along nodes at base of leaves. Leaves in whorls of 4, 
lanceolate-elliptic, 4.5-5.7 cm long, 1.2—1.5 mm broad, subobtuse or 
acute, narrowed at base into short petioles, thin, 3-veined, scabrous 
along margin due to very short, antrorse setae, elsewhere glabrous; 
floral leaves paired, uppermost leaves solitary, broadly elliptic or 
obovate, (1.5)3(5) mm long, (1)2.5(3.5) mm broad, with rounded apex, 
almost sessile, glabrous together with peduncles and pedicels. Panicles 
cuneate, broadly ovate, spreading, lax. Peduncles doubly furcately and 
ternately branched, 10—30-flowered. Corolla white, 2.5—4 mm across, 
with oblong-elliptic lobes. Fruits glabrous, rugose, 1 mm long, 1.5 mm 
broad. July to August. 

In open larch forests.—Soviet Far East: Ussuri, Uda River area. 
General distribution: North China (southeastern Manchuria). Described 
from southern part of Primorye (Vladivostok Region). Type in 
Leningrad. 

Note. In our territory, hybrids of this species with G. septentrionale 
Roem. et Schult. often occur, in which the mericarps usually are 
pubescent with scattered, sparse hairs or are pubescent in part and in 
the rest glabrous, but the pedicels are densely setose. 


Series 2. Borealia Pobed.—Mericarps smooth, pericarp adherent 
to seeds, usually densely setaceously hairy, rarely with scattered hairs 
or very rarely entirely glabrous. 


56. G. boreale L. Sp. pl. (1753) 108; Roem. and Schult. Syst. I, 
253; Ldb. Fl. alt. 1, 136, p.p.; DC. Prodr. IV, 600; Ldb. Fl. Ross. 2, 
412, Boiss. Fl. or. III, 48; Schmalh. Fl. II, 16; Grossh. Fl. Kavk. IV, 
30, p.p.; Kryl. Fl. Zap. Sib. X, 2581; The New Br. and Brown, 3, 
284.—G. trinervium Gilib. Fl. lith. I (1785) 12.—G. boreale var. 
kamtschaticum Maxim Prim. fl. amur. (1859) 141.—Iec.: Fedch. and 
Fler. Fl. 908; Syreistsch. Fl. Mosk. gub. III, 198; Rchb. Ic. Fl. Germ. 
XVII, 1186 (I).—Exs.: Fl. exs. austro-hung. No. 2202; Fl. exs. Reipubl.. 
Boh-Slov. No. 275; Fl. stir. No. 789; Pl. Finl. exs. No. 364. 

Perennial. Rootstock slender, woody, brown-red. Stems 1-6, 30— 
60 cm tall, stout, straight, rarely ascending, branched, rarely simple, 
scabrous along ribs due to rigid setae or short-pubescent and along 
edges (often at lower and upper internodes) on (rarely) all glabrous. 
Leaves linear or linear-lanceolate, (15)20(30) mm long, (2)3-4(5) mm 
broad, obtuse or subacute, 1-3 veined (lateral veins often indistinct), 
scabrous from stout, antrorse setae or softly and densely pubescent 
along veins beneath or along margin, sometimes also over entire surface 

350 or lower surface; floral leaves paired, ovate, (2)3—5(6) mm long, 15- 


B62 


2.5 mm broad, with short setae along margin and middle vein beneath 
or over entire lower surface. Panicle narrow, elongated, more or less 
compressed, not dense flowered or broad, spreading (in case of 
hybridization with species having similar type of panicle). Corolla 3- 
4 mm across, with ovate, acute lobes. Anthers yellow. Style bipartite 
from middle. Fruits pubescent with dense, or rarely scattered hooked 
white hairs, very rarely entirely glabrous. June to August. 

Meadows in scrub, forest edges, mountain slopes, riverbanks, thin, 
mixed forests and birch forests.—Arctic Region: Arctic Europe, Arctic 
Siberia, Chukotka, Anadyr. European USSR: Karelia-Lapland, Dvina- 
Pechora, Baltic Region, Ladoga-Ilmen, Upper Volga, Volga-Kama, 
Upper Dnieper, Middle Dnieper, Volga-Don, Volga Region, Upper 
Dniester, Black Sea Region (only rarely in northern part), Lower Don 
(only near Rostov, introduced), Lower Volga (near Stalingrad, 
introduced), Ural Region. Caucasus: Eastern and _ southern 
Transcaucasia. Western Siberia: All regions. Eastern Siberia: Yenisei, 
Lena-Kolyma, Soviet Central Asia: Aralo-Caspian (only in northern 
part, Turgai. Region). General distribution: Arctic Region, Scandinavia, 
Central and Atlantic Europe, Armenia-Kurdistan, Dzhungaria-Kashgaria, 
Mongolia (?). Described from northern Europe. Type in London. 

Note. Highly variable in all characters and requires critical 
examination and better understanding. 


57. G. amurense Pobed. sp. n. in Addenda XXII, 716.—G. boreale 
Q. genuinum Maxim. Prim. fl. amur. (1859) 141; Kom. Fl. Manchzh. 
III (1907) 499, p.p.—G. boreale 8. latifolium Turcz. Fl. baic.-dah. I 
(1845) 533, p.p. 

Perennial. Rootstock thick, short, brown-red, develops horizontal, 
slender shoots. Stem 30—80 cm tall, thick, straight with short, branches 
erect at tips, 4-angled, with poorly marked ribs only on lower 
internodes, glabrous, rarely puberulent, forming velutinous ring at 
nodes. Leaves in whorls of 4, broadly elliptic or oblong-lanceolate, 
(28)30-40(45) mm long, (6)10(25) mm broad, obtuse or gradually 
acuminate at apex, rounded at base, narrowed into very short velutinous 
petiole or sessile, 3—S-veined, glabrous on both surfaces, with fine 
scattered setae along margin; floral leaves paired, broadly ovate, (2)3— 
5(6) mm long, (1.5)2(3) mm broad, almost rounded at apex, glabrous. 
Inflorescence corymbose-paniculate, broad, often with almost 
horizontally divaricate branches. Peduncles and pedicels setulosely 
pubescent, rarely glabrous peduncles doubly ternate. Flowers as long 
as pedicels or 2 times as short. Corolla white, 5 mm across; lobes 
oblong, narrowed abruptly into a short, obtuse tip. Stamens with slender 
filaments; anthers yellow. Style bipartite in upper third. Fruits dark 


333 


351 brown, binate or with one mericarp, the other rudimentary, 1 mm 
long, 2 mm broad, covered with long, dense, straight, setaceous hairs. 
June to July. 

Riverbanks, meadows and forests.— Soviet Far East: Zeya-Bareya, 
Kamchatka. General distribution: Northern China. Described from 
Bureya River (village of Bakhareva). Type in Leningrad. 

Note. Within the geographica! range of this species, G. 
septentrionale Roem. et Schult. is widely distributed, giving rise to 
numerous hybrids with G. amurense. 


58. G. amblyophyllum Schrenk ex Fisch. et Mey. Enum. pl. nov. 
(1841) 56; Ldb. FI. Ross. II, 407; Fedch. Perech. rast. Turkest. III 
(1909) 140.—G. salsugineum Kryl. et Serg. in Tr. Biol. inst. Tomsk. 
univ. II (1936) 65.—G. boreale auct. fl. Sib. ex parte, non L. 

Perennial. Rootstock slender, horizontal, 1-3 mm thick, reddish. 
Stems from 1-6, 15-45 cm tall, ascending only at base, thereafter 
straight, slender, stout, short-pubescent, scabrous along ribs, due to 
acute, retrorse setae, in each node with shortened vegetative twigs. 
Leaves in whorls of 4, linear, with parallel revolute margins, 15-20 
mm long, 0.5-1.5, rarely 2 mm broad, obtuse, sessile, with one 
longitudinal vein, scabrous along margin, along vein beneath and 
sometimes on upper surface due to acute, scattered antrorse setae; 
floral leaves 1-2 mm long, 0.75—1 mm broad, elliptic, obtuse, sessile, 
beneath (sometimes on upper surface) with scattered appressed hairs. 
Inflorescence paniculate, elongated narrow, compressed. Peduncles and 
pedicels pubescent. Corolla 3—3.5 mm across, white with ovate, 
subacute lobes. Fruits 1-1.5-2 mm broad, glabrous or with dense, 
erect, short straight setae. June to August (Plate XIX, fig. 3). 

On solonetz, saline and wet meadows.— Western Siberia: Irtysh, 
Altai. Soviet Central Asia: Balkhash Region, Dzhungaria-Tarbagatai. 
Described from Balkhash Region. Type in Leningrad. 


59. G. turkestanicum Pobed. sp. n. in Addenda XXII, 717.—G. 
boreale auct. fl. As. med. non L. 

Perennial. Rootstock slender, creeping, reddish. Stems one-three, 
straight, simple or very weakly branched, sturdy, 30-55 cm tall, 
glabrous over entire length, only short-pubescent at nodes, lustrous or 
covered with very short hairs along ribs and edges on lower internodes, 
rarely short-pubescent over entire length. Leaves in whorls of 4, erect 
upward, in inflorescence opposite, linear or lanceolate-linear, 
(2.7)3.5(40) mm long, (3)4(9) mm broad, ratio of length to width 7— 
10, barely narrowly at apex, obtuse, rounded at base, with very short 
petiole, almost sessile, with 1-3 prominent veins, but lateral veins 


334 


never reaching apex, setosely scabrous along margin, elsewhere 
glabrous, rarely with sparse setae beneath along main vein or with 
scattered setae on upper surface. Panicle cuneate, more or less narrow; 
compressed at onset of anthesis, obovate and more or less spreading 
in fruit. Peduncles and pedicels glabrous or setose. Corolla 4-5 mm 
across, with 4 (sometimes 5) elliptic short-acuminate lobes. Fruits 
sparsely pubescent with hooked setae or glabrous (var. gymnocarpum 
m.). June to July. 

In alpine and subalpine meadows and on dry slopes in subalpine 
zone.— Soviet Central Asia: Dzhungaria-Tarbagatai, Pamiro-Alai, Tien 
Shan. General distribution: Dzhungaria-Kashgaria (Kuldzha). Described 
from Dzhungarian Ala Tau. Type in Leningrad. 

Note. This species is easily distinguished from the European G. 
boreale L. even in general appearance, owing to its longer and narrower 
leaves, which are usually erect and somewhat appressed to the stem, 
the weak branching of the stems, or the stems without branches, and 
the considerably larger inflorescence, sometimes reaching down as far 
as the middle of the stem. Forms gymnocarpum m. of this species, 
often with narrower leaves, is distributed over almost the entire 
geographical range of this species. 


60. G. mugodsharicum Pobed. sp. n. in Addenda XXII, 717.—G. 
boreale auct. Fl. As. med. non L. 

Perennial. Rootstock slender, funiform, brown-red, woody, rooting 
at nodes. Stems few, 10—35 cm tall, often branched from base of stem, 
straight, slender, stout with short dense pubescence over entire surface. 
Leaves in whorls of 4, ovate or lanceolate, (12)18(25) mm long, (3)6(7) 
mm broad, ratio of length to breadth 3-4, somewhat attenuated at 
apex, acute or obtuse, 3-veined, lateral veins usually not reaching the 
apex, along margin slightly recurved, stiff, pubescent with short 
appressed setae on both surfaces, sessile; floral leaves broadly ovate 
or elliptic, small, obtuse. Panicles terminal at apexes of stems and 
numerous lateral branches, almost to base of stem, forming lax, large 
inflorescence, equaling half the length of plant. Peduncles ternately or 
doubly ternately branched, many-flowered. Pedicels short, equaling 
flowers or 2 times as long, together with pedicels short-pubescent. 
Corolla white, 2.5—3 mm across; lobes elliptic, abruptly acuminate at 
tip, fimbrillate along margin or glabrous. Fruits binate, 1 mm long, 1- 
1.75 mm broad, diffusely pubescent with short hooked setae. June to 
July. 

On stony slopes of rivers, in rocks—European USSR: Volga-Kama 
(southern Urals). Western Siberia: Upper Tobol, Irtysh. Soviet Central 


353 


335 


Asia: Aralo-Caspian. Endemic. Described from Mugodzhary Hills. Type 
in Leningrad. 

Note. Differs from typical G. boreale L. in having numerous, 
considerably shorter stems, and in being highly pubescent over the 
entire plant and leaves on both surfaces. Especially close to this species 
are certain plants from Armenia and Georgia; however, we believe that 
these specimens belong to G. boreale, which undergoes some change 
under the xerophytic conditions of southern Transcaucasia. 


61. G. septentrionale Roem. et Schult. Syst. veg. III (1818) 253; 
DC. Prodr. IV, 601.—G. boreale auct. Pursh, Fl. Am. Sept. I (1814) 
104; Ldb. FI. Ross. II, 412, p.p.; Schmalh. Fl. II, 16, p.p.—G. boreale 
a. Ldb. FI. alt. I (1829) 136.—G. boreale var. intermedium Serg. in 
Tr. Biol. inst. Tomsk. univ. II (1936) 64, non DC.—G. boreale var. 
rubiifolium Serg. in Kryl. Fl. Zap. Sib. X (1939) 2581.—G. boreale 
B. latifolium Turcz. Fl. baic.-dah. I (1845) 533, p.p.—G. rubioides 
Schangin ex Pall. Neue nord. Beitr. VI (1793) 42. 

Perennial. Rootstock long, slender, reddish brown. Stems 30-80 
cm tall, single, rarely 2-6, straight, more rarely ascending, stout, simple 
or branched, glabrous, short-pubescent at nodes, lustrous, more rarely 
scabrous along ribs due to short stout setae; internodes long, 
considerably longer than leaves. Leaves in whorls of 4, lanceolate or 
oblong-lanceolate, (30)45-55(70) mm long, (4)7—8(12) mm broad, long 
attenuated at apex, acute or subobtuse, 3-veined, glabrous or scabrous 
along margin and veins beneath, very rarely with denser soft pubescence 
beneath; floral leaves lanceolate, 3-8 mm long, 2-3 mm broad. Panicle 
broad, spreading, terminal and in axils of 3-4 upper nodes, on long, 
glabrous or setulosely pubescent peduncles and pedicels, the latter 
equaling flowers or 2—3 times as long. Corolla white, 34 mm across, 
its lobes elliptic, abruptly short-acuminate. Fruits 1.25—1.5 mm long, 
1.5-—2 mm broad, pubescent with not very dense, hooked hairs. June to 
September. 

In taiga, deciduous forests, forests glades, scrub along river- 
banks.— Western Siberia: All regions. Eastern Siberia: Yenisei, Angara- 
Sayan, Dauria. Soviet Far East: Kamchatka, Zeya-Bureya. Soviet 
Central Asia: Aralo-Caspian, Balkhash Region, Dzhungaria-Tarbagatai, 
Tien Shan. General distribution: India-Himalayas, Dzhungaria- 
Kashgaria (Kuldzha), North America, Tibet. Described from North 
America. Type in Berlin. 

Note. This species varies, especially in the size and shape of the 
leaves. It is possible that a more detailed investigation will reveal 
narrower geographical races within the limits of the species. In typical 
plants, this species is well-distinguished from G. boreale L. by general 
appearance because of larger, broad leaves and a spreading panicle. 


354 


336 


However, given the variation of this species and the other one, the 
sharp differences between them disappear. 

The cytological studies of A. and D. Love (A. and D. Léve in 
Amer. Midl. Natur., 521 (1954) 97) showed that the European G. 
boreale L., is a tetraploid species (2n = 44), whereas the Asian- 
American G. septentrionale Roem. et Schult. is a hexaploid (2n = 66). 

Judging from several specimens of this species from America, 
their fruits are pubescent with straight hairs, whereas the Asian 
representatives of this species are covered with hooked hairs. 


Series 3. Valantioidea Pobed.—Cymes terminal on stems and 
branches forming short, wide, corymbose inflorescence; upper branches 
long, their corymbs almost reaching apical corymbs. 


62. G. valantioides M.B. FI. taur.-cauc. I (1808) 102; DC. Prodr. 
IV, 599; Ldb. Fl. Ross. I, 411; Boiss. Fl. or. III, 49; Grossh. FI. 
Kavk. IV, 30.—Exs.: GRF, No. 269. 

Perennial. Stems one or several, highly branched, 15-35 cm tall, 
straight; lower internodes terete, upper 4-angled, smooth, glabrous, 
rarely densely pubescent with short, erect hairs. Leaves in whorls of 
4, rhombic-lanceolate, (11)20-30(50) mm long, (5)8(14) mm broad, 
3-veined; floral leaves considerably smaller, attenuated at apex, 
subobtuse, scabrous along veins on both surfaces and margin, rarely 
hairy over entire surface beneath, with short petiole. Inflorescence 
corymbose. Peduncles doubly ternately and in the latter branched 
furcately, apical and lateral branches many-flowered, straight, glabrous 
or pubescent. Pedicels short, equaling flowers or 2—3 times as long. 
Corolla 2.5-3 mm across, white, brownish when dry, with elliptic, 
short-acuminate lobes, with incurved tip. Fruits glabrous, tuberculate, 
tiny, 1-1.75 mm long, 1.5—2.5 mm broad. June to August (Plate XIX, 
fig. 4). 

On rocks, in subalpine and alpine zone, in meadows.— Caucasus: 
All regions, except southern Transcaucasia. Described from Kislovodsk. 
Type in Leningrad. 


Section 10. XANTHOGALIA DC. Prodr. 4 (1830) 603 pro subsect.— 
Chromogalia Boiss. FI. or. III, 60.—Panicles terminal, many-flowered. 
Flowers bisexual, yellow, rarely cream colored. Leaves in whorls of 
6-14, single-veined, with cusp at apex. Fruits pubescent, with hairs 
not curved at tip, or glabrous. Perennial herbs with straight, stout, 
glabrous, or pubescent stems. 


S5i 


337 


Series 1. Vera Pobed.—Fruits glabrous, very rarely pubescent. 
Leaves not reflexed at anthesis. Anthers yellow. 


63. G. verum L. Sp. pl. (1753) 107; M.B. FI. taur.-cauc. 1, 103; 
DC. Prodr. IV, 603, p.p.; Ldb. Fl. Ross. III, 414, p.p.; Boiss. Fl. or. 
III, 62, p.p.; Schmalh. FI. II, 14; Grossh. Fl. Kavk. IV, 31; Kryl. FI. 
Zap. Sib. X, 2585, p.p.—G. luteum Moench. Meth. pl. (1794) 485; 
Gilib. Fl. lith. I (1781) 9.—G. verum var. leiophyllum Wallr. Sched. 
crit. (1822) 56.—G. verum var. trachyphyllum Wallr. 1.c.—G. verum 
f. glabrum Rgl. et Herd. Enum. pl. Sem. 3 (1868) 9.—G. berum f. 
puberulum Serg. in Tr. Biol. inst. Tomsk. univ. (1936) 68.—G. verum 
f. hispidulum Rgl. et Herd. l.c.—G. verum L. var. Trin. in herb.—G. 
minutum L. Sp. pl. (1753) 154.—Ic.: Syreistsch. Fl. Mosk. gub. III, 
196; Fedch. and Fler. Fl. 910; Rchb. Ic. Fl. Germ. XVII, tab. 1187.— 
Exs.: Fl. Boh. and Morav. exs. No. 1089; Fl. Pol. exs. No. 346; PI. 
Finl. exs. No. 365; Fl. Gall. and Germ. exs. No. 1494. 

Perennial. Rootstock long, slender, creeping, short, thickened, 
branched. Stems one-several, 30-125 cm tall, 4-angled, straight, simple 
or with short, sterile shoots, glabrous or glabrous below with middle 
and upper internodes more or less densely pubescent or pubescent 
only under nodes and glabrous elsewhere. Leaves in whorls of 6—10— 
15, more or less approximate or distant, linear or filiform, (10)18— 
20(35) mm long, (0.5)—-1(1.5) mm broad, usually strongly revolute 
along margin, almost up to midrib or oblong-lanceolate, with barely 
revolute margin (var. sibiricum m.), with short cusp at apex, narrowed 
at base, upper surface dark green, lustrous, glabrous or with short, 
acute, antrorse setae or with similar setae only along margin, beneath 
light green, glabrous; floral leaves in whorls of 4-6; uppermost paired 
or solitary, very small, 3-5 mm long, 0.25—0.5 mm broad. Peduncles 
doubly ternately branched, many-flowered. Inflorescence paniculate, 
dense or broadly spreading (var. sibiricum m.) terminal on stems and 
upper branches, together with pedicels glabrous or scabrous due to 
acute, short setae. Corolla bright yellow, with honey odor, 3 mm across, 
with oblong, subobtuse lobes. Anthers yellow. Style bipartite from 
middle or slightly above. Fruits binate, rarely one mericarp 
underdeveloped, 1.5 mm long, glabrous or pubescent (var. sibiricum), 
finely unevenly tuberculate. June to September. 

Steppes, steppe slopes, edges and glades in deciduous forests and 
in upland meadows.—Arctic Region: Chukotka (introduced). European 
USSR: Karelia-Lapland, Dvina-Pechora, Baltic Region, Ladoga-IImen, 
Upper Volga, Volga-Kama, Upper Dnieper, Middle Dnieper, Volga- 
Don, Black Sea Region, Crimea, Lower Don, Lower Volga. Caucasus: 
Ciscaucasia, Dagestan, western, eastern and southern Transcaucasia, 


338 


Plate XX. 

355 1. Galium subuliferum Somm. and Lev., upper part of plant, corolla, fruit— 
2. G. bullatum Lipsky., upper part of plant, corolla, ovary with styles, fruit 
with one dehisced mericarp.— 3. G. majmechense Bordz., upper part of plant, 
corolla, fruit— 4. G. brachyphyllum Roem. and Schult., habit, corolla, fruit.— 

5. G. kutzingii Boiss., upper part of plant, leaf whorl, fruit, corolla. 


358 


339 


Western Siberia: All regions. Eastern Siberia: All regions. Soviet 
Central Asia: Tien Shan, Dzhungaria-Tarbagatai. General distribution: 
Scandinavia, Central and Atlantic Europe, western and eastern 
Mediterranean, Balkans-Asia Minor, North America (introduced). 
Described from Western Europe. Type in London. 

Note. This variable species often hybridizes with species of other 
sections. Siberian plants of the species differ somewhat from European 
ones, having leaves oblong-linear, slightly longer with barely revolute 
margins and always glabrous on both sides; panicle more spreading, 
broad; and stem taller, glabrous. In Eastern Siberia and in the Soviet 
Far East, the widespread form has pubescent fruit (var. sibiricum m.). 
In Western Siberia, where it is sometimes found with G. ruthenicum, 
hybrids: between these two occur some with a pubescent stem, others 
with pubescent leaves. G. verum var. sibiricum extends as far west as 
Dvina-Pechora and Karelia-Lapland, where it hybridizes with G. 
ruthenicum. This variety could be considered a separate geographical 
race, if the form with glabrous fruits were not distributed over the 
entire geographical range, the latter being indistinguishable from it in 
other characteristics and having a resemblance to typical European G. 
verum L. 


64. G. subuliferum Somm. and Lev. in Tr. SPb. bot. sada, XII 
(1892) 151 and XVI (1900) 206; Grossh. Fl. Kavk. IV, 33 and Opred. 
rast. Kavk. 253. 

Perennial. Root long, thicker, dark reddish brown. Stems several, 
ascending, woody at base, thickened at nodes, 30-50 mm tall, 4-angled, 
with white hairs along ribs, light, lustrous, glabrous, branched, with 
more or less long branches erect at tip. Leaves in whorls of 6, filiform, 
very narrow, (8)10(18) mm long, 0.25—0.75 mm broad, with white 
cusp at apex and one white vein sharply projecting below, abruptly 
revolute along margin, thicker, sessile, glabrous on both surfaces, 
scabrous along margin due to short, rigid, antrorse setae, upper leaves 
considerably smaller; floral leaves paired or solitary. Inflorescence 
paniculate, long, occupying most parts of stem, lax, with branches 
erect at tip. Peduncles ternately branched, elongated, slender, filiform. 
Pedicels 2 times as long as flowers; peduncles and pedicels glabrous. 
Flowers yellow, 3 mm across excluding length of cusps; lobes ovate, 
with long acuminate tip (equaling half the length of lobes). Stamens 
with slender filament gradually broadened at base; anthers large, brown. 
Style bipartite up to one third. Fruits binate, tiny, 1-1.25 mm long, 
1.5—2 mm broad, glabrous, smooth. June to August (Plate XX, fig. 7). 


359 


340 


On dry, open, sunny slopes in rocks and stony cliffs.—Caucasus: 
Western, eastern and southern Transcaucasia. Endemic. Described from 
Adzhar Range (between Khula and Shvakheri). Type in Leningrad. 


65. G. wirtgenii F. Schultz, Arch. de Fl. I (1854-1855) 201; 
Lindm. Svensk fanerog. 508.—G. verum var. wirtgenii Abrom. FI. 
Ost-u. Westpreuss. (1898-1940) 360; Szaf. Kulcz. and Pavl. Résliny 
Polsk. (1953) 618.—G. verum B. praecox Lang. in Hagenbach, FI. 
Basil. Suppl. (1843) 26.—G. verum ssp. praecox Petrak in Allgem. 
Bot. Zeitschr. Jahrg. XVI, 2 (1910) 21; Hegi, Ill. Fl. VI, 1.—G. praecox 
(Lang) Petrak in Résliny Polsk. (1924) 572.—G. verum auct.—Exs.: 
Fl. exs. austro-hung. No. 2222 (sub G. praecox Lang.); Fl. siles, exs. 
No. 826 (sub G. praecox Lang.); Fl. Boh. and Morav. exs. No. 1088. 

Perennial. Rootstock creeping, slender, reddish brown. Stems 1-— 
10, stout, straight, 45-75 cm tall, simple or with short, vegetative 
shoots, glabrous on lower internodes, short-pubescent elsewhere. 
Leaves in distinct whorls of 6-10, oblong-linear, (18)25(30) mm long, 
0.5(1)2 mm broad, at tip with short cusp at apex, with slightly revolute 
margin, glabrous or scabrous along margin and sometimes on upper 
surface due to short antrorse setae; floral leaves in whorls of 4—6, 
smaller, uppermost paired or solitary, elliptic or lanceolate, 1.5—3 mm 
long, 0.25-1 mm broad. Inflorescence paniculate, interrupted, few- 
flowered, its lateral branches always shorter than adjacent internode, 
reaching, up to or almost middle of stem. Peduncles usually short- 
pubescent. Pedicels glabrous, reflexed in fruit, Corolla golden yellow, 
odorless, 3 mm across, with 4—5 ovate, subobtuse lobes. Anthers yellow. 
Style bipartite above middle. Fruits binate, 1 mm long, 1.5 mm broad, 
glabrous, dark brown, almost dark, finely tuberculate. May to June. 

In meadows, forest edges, on sandy and sandy-clayey soils and 
limestones.—European USSR: Baltic Region, Upper Dniester, 
Bessarabia. General distribution: Central Europe, North America 
(introduced). Described from the Rhine. Type in Vienna. 

Note. This species is well distinguished from G. verum by the 
widely spaced whorls of leaves with almost nonrevolute leaf-blade 
margins; lax, interrupted inflorescence, reaching almost to the middle 
of the stem; odorless flowers; and earlier flowering, preceding the 
flowering of G. verum by almost a month. The first to draw attention 
to the distinctiveness of this species was Lang who also named it in 
1843 as variety praecox of G. verum L. It is interesting to note that in 
exsiccatae (Fl. austro-hung., Fl. siles), it is issued under the name G. 
praecox Lang, although it is never called such in the literature. Wirtgen, 
who collected it in the Rhine Valley, also referred it to the different 
species G. eminens Gren. et Godr., although incorrectly, a fact also 


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noted by Schultz [Arch. de fl. (1855) 192], who proposed for the plant 
from the Rhine a new specific name.—G. wirtgenii. 


66. G. fistulosum Somm. et. Lév. in Bull. Soc. Bot. Ital. ser. II, 
III (1894) 31; Tr. SPb. bot. sada, XIII, 195 and XVI, 209; Grossh. FI. 
Kavk. IV, 31.—Ic.: Somm. and Lév. in Tr. SPb. bot. sada, XVI, tabl. 
XXII, fig. 1-7. . 

Perennial. Rootstock slender, horizontal, long. Stems numerous, 
7-25 cm tall, woody at base, ascending, fistular, pale green, lustrous, 
4-angled, with sharply projecting almost winglike ribs, glabrous or 
short curly pubescence over entire surface or only at nodes. Leaves in 
whorls of 8—9, more or less approximate, lanceolate-linear, (6)7—8(15) 
mm long, (0.5)0.75—1.5(2) mm broad, slightly or strongly revolute 
along margin, narrowed toward base, apex cuspidate, both surfaces 
glabrous, upper surface blackened on drying, beneath light green, 
single-veined, leaves in whorls of 4—6, in inflorescence uppermost 
floral leaves paired or solitary. Panicles narrow, elongated, interrupted. 
Peduncles doubly branched, short, few-flowered. Pedicels equaling 
flowers, pubescent together with peduncles. Corolla yellow, 2—3.5 mm 
across, with obtuse, oblong lobes, scabrous outside. Anthers yellow. 
Style bipartite almost from base. Fruits smooth, glabrous. July to 
August. 

Along rubbly slopes and screes in alpine zone.— Caucasus: Western 
Transcaucasia kenky northern part). Described from Teberda Range. 
Type lost? 


67. G. anfractum Somm. et, Lév. in Bull. Soc. Bot. Ital. ser. II, 
III (1894) 29; Grossh. Fl. Kavk. IV, 31.—G. verum auct. fl. Cauc. 
non L. 

Perennial. Rootstock slender, incites at Stems 1-10, 20-70 cm 
tall, stout, straight, simple or with few short vegetative shoots, 
pubescent with short hairs over entire surface, sometimes only under 
nodes or, rarely glabrous on lower internodes, above short-pubescent. 
Leaves in more or less distant whorls of 8-10, oblong-linear, (9)20— 
25(35) mm long, (0.5)1(1.5) mm broad, with barely revolute, rarely 
strongly revolute margin, with long cusp at apex, upper surface dark 
green, beneath light green, with whitish, broad, prominent midrib, 
glabrous or scabrous on upper surface or only along margin, due to 
short antrorse setae, lower floral leaves in whorls of 4—6, upper ones 
paired, uppermost solitary, (1.5)2(4) mm long, 0.5—0.75 mm broad, 
elliptic or oblong-elliptic, with long cusp at apex. Inflorescence 
paniculate, spreading, lax, interrupted, often absent below middle of 
stem, with more or less short, lateral branches. Peduncles doubly 


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ternately branched, densely short-pubescent together with pedicels. 
Pedicels equaling flowers or somewhat longer, declinate in fruit. Corolla 
yellow, large, 4 mm across with elliptic or oblong-elliptic lobes, obtuse 
or short acuminate and in latter case with incurved tip. Anthers yellow. 
Style bipartite almost from middle. Fruits with one mericarp, another 
underdeveloped, rarely binate, 1 mm long, 2 mm broad, glabrous, 
tuberculate. 

In alpine and subalpine meadows, on moraines and screes.— 
Caucasus: Ciscaucasia, Dagestan, western, eastern and southern 
Transcaucasia. General distribution: Armenia-Kurdistan. Described 
from Utviry Range (Svanetia). Type in Leningrad. 


68. G. majmechense Bordz. in Vestn. Kievsk. bot. sada, V—VI 
(1927) 21.—G. verum B. consanguineum Boiss. FI. or. III (1875) 62.— 
G. sanguineum auct. non Boiss.: Grossh. Fl. Kavk. IV, 31.—Exs: PI. 
or. exs. No. 224 (sub G. consanguineum Boiss.). 

Perennial. Root thick, woody. Stems numerous, 30-60 cm tall, 
somewhat ascending, weakly branched, subterete, with smooth ribs, 
glabrous, with ring or velvety pubescence at base of leaves or rarely 
short hooked pubescence; lustrous. Leaves in whorls of 8—12, oblong- 
linear, (20)25(28) mm long, (1)2(3) mm broad, with thick stout cusp, 
weakly revolute and scabrous along margin due to rigid, antrorse setae, 
elsewhere glabrous, sessile, upper surface lustrous, blackened on drying, 
pale green beneath; floral leaves in whorls of 3-6, uppermost leaves 
paired or solitary, much smaller than cauline leaves. Inflorescence 
paniculate, terminal and in axils of upper leaves, more or less narrow, 
oblong, not very dense, many-flowered. Peduncles doubly ternate, short. 
Pedicels equaling flowers or shorter, glabrous. Corolla yellow, 3 mm 
across with ovate lobes, having thick short mucro at tip. Anthers orange- 
yellow, brownish yellow on drying. Stigma globose. Fruits 1.5 mm 
long, with one mericarp, 1 mm broad or binate, glabrous. July to August 
(Plate XX, fig. 3). 

Meadows in alpine zone—Caucasus: Eastern and southern 
Transcaucasia. General distribution: Armenia-Kurdistan, Iran. 
Described from Armenia (Maimek Mountains). Type in Kiev. 

Note. This species differs from G. consanguineum Boiss., described 
from southern Iran, by the large number of leaves in the whorls, 8-12 
(and not 6), larger flowers, and also narrower leaves and a more 
spreading panicle. 


69. G. pamiro-alaicum Pobed. sp. n. Addenda XXII, 718.—G. 
verum auct. fl. As. med. non L. 


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Perennial. Root thick, vertical, reddish brown. Stems 2-10 in 
number, 40-100 cm tall, completely glabrous, even in inflorescence 
or short-pubescent over entire length, or glabrous below and pubescent 
upwards, or pubescent below and glabrous upwards and in inflorescence 
straight, stout, lustrous, subterete below, 4-angled above. Leaves in 
distant whorls of (6)8(10), oblong-linear, (20)40(50) mm long, 
(0.5)1(2.5) mm broad, narrowed at base, sessile, with cusp at apex, all 
or only lower ones with barely revolute margin, then the remaining 
with strongly revolute margin up to midrib, single-veined, glabrous 
on both surfaces or with short, white, soft hairs beneath, upper surface 
glabrous or scabrous due to short, rigid setae; floral leaves in whorls 
of 6, smaller, becoming shorter toward tip of inflorescence, reflexed 
at anthesis in branches of inflorescence, paired or solitary, elliptic, 
(1.5)2(3) mm long, (0.25)0.5(1) mm broad, together with peduncles 
and pedicels glabrous, rarely pubescent. Pedicels 2—3 times as long as 
fruits. Inflorescence paniculate, long, equaling third or half the length 
of stem, spreading, not dense, at bottom supplemented with paniculate 
inflorescences at ends of axillary branches, often hanging below middle 
of stems. Corolla yellow, 3.5—4 mm across, with oblong or ovate 
lobes. Anthers oblong, yellow. Style bipartite somewhat above middle. 
Fruits arcuate, binate or with one rudimentary mericarp, 1 mm long, 
2 mm broad, glabrous, brownish. May to September. 

On stony and steppe mountain slopes, in subalpine meadows and 
in middle-mountain belt, on screes, in scrub, open maple forests.— 
Soviet Central Asia: Balkhash Region, Dzhungaria-Tarbagatai, 
mountainous Turkmenia, Syr-Darya, Pamiro-Alai, Tien Shan. 
Endemic. Described from Gardan-Ushti Range in Pamiro-Alai. Type 
in Leningrad. 


Series 2. Aurea Pobed.—Fruits glabrous. Leaves reflexed at 
anthesis. Anthers black. 


70. G. aureum Visiani in Orto Bot. Padov. (1842) 134, in not.; 
Boiss. Fl. or. III, 61; Grossh. Fl. Kavk. IV, 31.—Iec.: Rchb. Ic. Germ. 
XVII, tab. 1188. 

Perennial. Entire plant light green. Rootstock vertical, woody, not 
thick, reddish brown. Stems 1-2, 30-60 cm tall, woody at base, 
branched or simple, ascending, 4-angled, with sharply prominent, white 
bands along ribs, glabrous, rarely with patent, scattered hairs over 
entire surface or only under nodes. Leaves in more or less distant 
whorls of 6-10, oblanceolate, 8(10)—20(25) mm long, (1.5)2.5(4) mm 
broad, obtuse, with more or less long cusp, gradually long attenuated 
toward base, slightly revolute, scabrous along margin due to short, 


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rigid setae or glabrous, for the rest glabrous on both surfaces; floral 
leaves in whorls of 6, uppermost leaves paired or solitary, all 
considerably smaller than cauline leaves, gradually reduced toward tip 
of stem, (1)2(2.5) mm long, (0.25)0.5(0.75) mm broad. Inflorescence 
paniculate, oblong narrow, with upright or horizontally inclined, short 
branches. Peduncles few-flowered, short. Pedicels equaling flowers in 
fruit 2 times as long as fruits, pedicels and peduncles glabrous or 
rarely with scattered, patent hairs. Corolla golden yellow, 3.5-4 mm 
across, with oblong lobes having long cusp at tip. Anthers black. 
Style bipartite below middle. Fruits binate or with one rudimentary 
mericarp, 2 mm long, 2.5 mm broad, glabrous, yellow, finely 
tuberculate. May to August. 

On limestone, slopes of mountains, stony riverbanks, subalpine 
meadows, in rocks.— Caucasus: Ciscaucasia, Dagestan, western 
Transcaucasia (only northern part), eastern Transcaucasia. General 
distribution: Western Mediterranean (only northern Italy), Balkans- 
Asia Minor (?) (specimens from this area differ slightly from the 
typical). Described from Padua. Type in Italy. 


71. G. achurense Grossh. in Bot. mat. Gerb. Bot. inst. AN SSSR, 
XIII (1950) 25. 

Perennial. Rootstock slender, reddish brown. Stems 7-15, 25-30 
cm tall, simple, branched towards tip, indistinctly 4-angled, along ribs 
setulosely scabrous. Leaves in more or less approximate whorls of 6— 
10, oblong-linear, 7-8 mm long, 1-1.5 mm broad, with barely revolute, 
smooth margin, short-acuminate at apex, barely narrowed toward base, 
with one weakly prominent midrib, glabrous on both surfaces, reflexed 
at anthesis; floral leaves in whorls of 6, considerably smaller than 
cauline leaves, uppermost paired or solitary, ovate, 1.5 mm long, 1 
mm broad. Inflorescence paniculate, spreading, cymes few-flowered. 
Peduncles and pedicels glabrous, the latter perpendicular to peduncle 
bearing it, branched cymes often with rudimentary flowers in form of 
lilac buds. Corolla milk white, 3 mm across, with ovate, obtuse lobes. 
Anthers black. Style bipartite from middle, at base surrounded by 
fleshy discoid nectary. Young fruits glabrous. May. 

On stony slopes and along shingles.—Caucasus: Southern 
Transcaucasia. Endemic. Described from village of Akhur Nakhichevan 
ASSR. Type in Leningrad. 


Series 3. Ruthenica Pobed.—Fruits pubescent with straight hairs, 


_ very rarely glabrous. Leaves not reflexed at anthesis. Anthers yellow 


or brown. 


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72. G. lacteum (Maxim.) Pobed. sp. n. in Addenda XXII, 718.— 
G. verum var. lactiflorum Maxim. in sched.—G. verum var. lacteum 
Maxim. in Mél. Biol. IX (1873) 265.—G. verum auct. fl. Or. extr. non 
L.: Thunbg. FI. Jap. (1784) 59; Kom. and Alis. Opred. rast. Dalnevost. 
kr. II, 948. | 

Perennial. Rootstock thick, horizontal, reddish brown. Stems 1— 
10, stout,. straight, simple or with few branches at base, 40-100 cm 
tall, below subterete, glabrous, higher up 4-angled, with short or velvety 
pubescence, pubescent with fairly long, dense, patent hairs in upper 
internodes and in inflorescence or densely patently pubescent 
throughout. Leaves in distant whorls of 8-14, all or only middle ones 
oblong-linear, remaining filiform linear, (20)35-45(55) mm long, 
(0.5)0.75(2.5) mm broad, with extremely short mucro at apex, gradually 
narrowed at base, sessile, along margin slightly revolute or strongly 
revolute up to midrib, whitish beneath due to dense, short pubescence, 
upper surface glabrous or scabrous due to scattered, short, antrorse 
setae or, rarely, glabrous on both surfaces; floral leaves in whorls of 
5-6, becoming smaller toward tip of stem, uppermost leaves paired or 
solitary, elliptic or oblong-elliptic, often patently pubescent on both 
surfaces, (1.5)2(4) mm long, (0.25)0.5(1) mm broad. Inflorescence 
paniculate, oblong, more or less spreading. Cymes fairly densely 
aggregated on branches of panicle. Peduncles and pedicels densely 
patently pubescent, the latter equaling flowers or shorter. Corolla cream 
colored, 4-lobed, rarely 5-lobed, 2.5—3.5 mm across, with oblong, short 
acuminate lobes. Anthers brown, darkened on drying. Style bifid above 
middle. Fruits binate, 1 mm long, 2 mm broad. Mericarps crescent 
shaped, covered with straight, erect hairs. June to September. 

In meadows, along riverbanks, in scrub, in mixed forests.—Soviet 
Far East: Ussuri. General distribution: Japan, China. Described from 
Primorsky Province (Posjet Bay). Type in Leningrad. 

Note. Galium lacteum is close to G. ruthenicum, but well 
distinguished from it because of considerably longer leaves, cream 
colored flowers and longer, spreading pubescence in the inflorescence. 

C.J. Maximovicz indicates that in those places where var. lacteum 
with cream-colored flowers grows, the plant with yellow flowers is 
also seen. It is difficult to judge the color of flowers in the herbarium; 
however, G. lacteum in the Primorsky province is extremely consistent, 
and specimens with yellow flowers are G. verum var. sibiricum Pobed. 


73. G. densiflorum Ldb. FI. alt. I (1829) 137 and FI. Ross. III, 
413; Kryl. Fl. Zap. Sib. X, 2583.—Ic.: Ldb. Ic. pl. Fl. Ross. II, tab. 
194. 


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Perennial. Rootstock horizontal, not thick, reddish brown. Stems 
1-3, 40-70 cm tall, subterete, above 4-angled, straight, stout, simple 
or weakly branched at base, lower internodes glabrous, middle often 
short pubescent sometimes velvety pubescent, often diffusely patently 
pubescent in inflorescence. Leaves in distant whorls of 8, oblong- 
linear, (3.5)4(4.5) cm long, (0.75)1.5(3) mm broad, with cusp at apex, 
gradually narrowed at base, with weakly revolute margin, single-veined, 
white-green beneath, dark green on upper surface, both surfaces 
glabrous; floral leaves in whorls of 6, differing only a little in length 
from cauline leaves, in twos or solitary in branches of inflorescence, 
oblong-elliptic, (2)2.5(4) mm long, (0.25)0.5(0.75) mm broad, upper 
surface glabrous, pubescent beneath or glabrous on both surfaces, with 
mucro at apex. Inflorescence paniculate, long, equaling a third or the 
length of stem or somewhat longer, compressed. Peduncles and pedicels 
diffusely patently pubescent, or pedicel glabrous. Corolla yellow, 3.5 
mm across, with elliptic mucronate lobes. Anthers light brown. Style 
bifid, somewhat above the middle. Fruits often with one mericarp, the 
other rudimentary, often many fruits completely undeveloped, 
mericarps 1.75 mm long, 1 mm broad, weakly curved, covered with 
straight, erect hairs. July to August. 

In lower zones of alpine belt, on open stony slopes, in wet meadows, 
fields, along riverbanks.—European USSR: Karelia-Lapland, Dvina- 
Pechora. Western Siberia: Ob’ Region, Altai. Eastern Siberia: Yenisei, 
Lena-Kolyma, Angara-Sayan, Dauriya. Described from Zmeinogorsk. 
Type in Leningrad. 


74. G. ruthenicum Willd. Sp. pl. I (1797) 597; M.B. FI. taur.- 
cauc. III, 108; Ldb. Fl. alt. I. 138; Grossh. Fl. Kavk. IV, 32.—G. 
verum Y. trachycarpum DC. Prodr. IV (1830) 603.—G. verum B. 
lasiocarpum Ldb. Fl. Ross. III (1844-1846) 415.—G. verum f. 
subpubescens Serg. in Tr. Biol. inst. Tomsk. univ. (1936) 68.—G. 
verum var. tomentosum Rgl. and Herd. Enum. pl. Sem. (1864) 10.— 
Exs.: GRF, No. 1669; Pl. Finl. exs. No. 366, 957; Dorfler. Herb. 
norm. No. 4529; Fl. Pol. exs. No. 45. 

Perennial. Rootstock slender, creeping, reddish brown. Stems one— 
numerous, 25-30 cm tall, straight, stout, simple or branched, densely 
short pubescent